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<document ID-DOI="10.7717/peerj.9192" ID-Zenodo-Dep="5824972" approvalRequired="3" approvalRequired_for_document="2" approvalRequired_for_treatments="10" checkinTime="1641472430154" checkinUser="jeremy" docAuthor="Carr, Thomas D." docDate="2020" docId="038A657CFFC8FFF5FE14FD7F3415F96D" docLanguage="en" docName="Carr2020.pdf.imf" docOrigin="PeerJ 8" docStyle="DocumentStyle:B6346867C9D49AB2DD3A1D607021D2E9:PeerJ.2019-.journal_article" docStyleId="B6346867C9D49AB2DD3A1D607021D2E9" docStyleName="PeerJ.2019-.journal_article" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="2" lastPageNumber="94" masterDocId="FFB31D04FFC4FFA8FFE6FF88371AFFBC" masterDocTitle="A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence" masterLastPageNumber="e 9192" masterPageNumber="e 9192" pageNumber="13" updateTime="1673632140161" updateUser="felipe">
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<mods:titleInfo>
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<mods:title>A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Carr, Thomas D.</mods:namePart>
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<mods:affiliation>Department of Biology, Carthage College, Kenosha, WI, USA</mods:affiliation>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>PeerJ</mods:title>
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<mods:part>
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<mods:date>2020</mods:date>
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<mods:detail type="volume">
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<mods:number>8</mods:number>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.7717/peerj.9192</mods:identifier>
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<mods:identifier type="Zenodo-Dep">5824972</mods:identifier>
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<treatment LSID="urn:lsid:plazi:treatment:038A657CFFC8FFF5FE14FD7F3415F96D" httpUri="http://treatment.plazi.org/id/038A657CFFC8FFF5FE14FD7F3415F96D" lastPageId="93" lastPageNumber="94" pageId="12" pageNumber="13">
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<subSubSection lastPageId="61" lastPageNumber="62" pageId="12" pageNumber="13" type="description">
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<paragraph blockId="12.[498,1542,759,1907]" box="[498,772,759,789]" pageId="12" pageNumber="13">
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<heading bold="true" box="[498,772,759,789]" fontSize="12" level="2" pageId="12" pageNumber="13" reason="0">
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<emphasis bold="true" box="[498,772,759,789]" pageId="12" pageNumber="13">Cladistic analyses</emphasis>
|
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</heading>
|
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</paragraph>
|
||
<paragraph blockId="12.[498,1542,759,1907]" pageId="12" pageNumber="13">
|
||
The reduced cladistic analysis of 31 specimens, following the New Technology and Traditional searches, recovered one most parsimonious tree (i.e., ontogram) of 3,053 steps, with an ensemble Consistency Index (CI) excluding uninformative characters of 0.65, an ensemble Homoplasy Index (HI) of 0.35, an ensemble Retention Index (RI) of 0.72, and an ensemble Rescaled Consistency Index (RCI) of 0.50. The ontogram recovered 21 growth stages; in the adult region of the ontogram several branches contain multiple specimens; these are most simply interpreted as specimens of the same maturity (
|
||
<figureCitation box="[1460,1525,1044,1070]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="12" pageNumber="13">Fig. 2</figureCitation>
|
||
). Bremer, jackknife, and bootstrap values are shown in
|
||
<figureCitation box="[1134,1216,1083,1110]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="12" pageNumber="13">Fig. 1B</figureCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="12.[498,1542,759,1907]" pageId="12" pageNumber="13">
|
||
The ontogram is composed of 21 growth stages, including the group of most mature specimens, and all but the first and third are supported by unambiguously optimized synontomorphies (Data S4). Despite the addition of 26 specimens and 1,766 characters, these results are congruent with those obtained by
|
||
<bibRefCitation author="Carr TD & Williamson TE" box="[1100,1410,1243,1269]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="12" pageNumber="13" pagination="479 - 523" part="142" refId="ref52732" refString="Carr TD, Williamson TE. 2004. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142 (4): 479 - 523 DOI 10.1111 / j. 1096 - 3642.2004.00130. x." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">
|
||
<emphasis box="[1100,1410,1243,1269]" italics="true" pageId="12" pageNumber="13">Carr & Williamson (2004)</emphasis>
|
||
</bibRefCitation>
|
||
, where
|
||
<materialsCitation box="[498,660,1283,1309]" collectionCode="LACM" pageId="12" pageNumber="13" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
,
|
||
<materialsCitation box="[673,828,1283,1310]" collectionCode="CMNH" pageId="12" pageNumber="13" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
,
|
||
<materialsCitation box="[841,1003,1283,1310]" collectionCode="LACM" pageId="12" pageNumber="13" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1016,1251,1283,1310]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="12" pageNumber="13" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
, and
|
||
<materialsCitation box="[1317,1476,1283,1309]" collectionCode="LACM" pageId="12" pageNumber="13" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
were again recovered at progressively mature growth stages (
|
||
<figureCitation box="[1153,1246,1323,1349]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="12" pageNumber="13">Figs. 1B</figureCitation>
|
||
and
|
||
<figureCitation box="[1307,1321,1323,1349]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="12" pageNumber="13">2</figureCitation>
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||
).
|
||
</paragraph>
|
||
<paragraph blockId="12.[498,1542,759,1907]" pageId="12" pageNumber="13">
|
||
The a posteriori analysis that included an artificial adult resulted in seven 3,071-step MPTs that, in a strict consensus ontogram did not group with a single, presumably most mature, specimen. Therefore, a different approach was taken, specifically the greatest distance from the root. Of that group,
|
||
<materialsCitation box="[955,1140,1482,1509]" collectionCode="FMNH" pageId="12" pageNumber="13" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
possessed the greatest number of autontomorphies, 19 character changes, in contrast to the five others; ergo, that specimen was regarded as the most mature of the group, having undergone the greatest amount of change, and so represents the twenty-first growth stage (i.e., the terminus of the growth series). In contrast, the massive specimen
|
||
<materialsCitation box="[994,1139,1641,1668]" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
, previously regarded as the most mature individual
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[779,846,1682,1707]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[779,846,1682,1707]" italics="true" pageId="12" pageNumber="13">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[864,1243,1681,1708]" journalOrPublisher="Anatomical Record" pageId="12" pageNumber="13" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[864,1243,1681,1708]" italics="true" pageId="12" pageNumber="13">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
), was recovered as one of the least mature adults.
|
||
</paragraph>
|
||
<paragraph blockId="12.[498,1542,759,1907]" lastBlockId="14.[498,1542,780,1444]" lastPageId="14" lastPageNumber="15" pageId="12" pageNumber="13">
|
||
The two purported female specimens (
|
||
<materialsCitation box="[976,1161,1761,1787]" collectionCode="BMRP" pageId="12" pageNumber="13" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1172,1305,1761,1788]" collectionCode="MOR" pageId="12" pageNumber="13" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
), a sex identification based on the presence of femoral medullary bone (
|
||
<bibRefCitation author="Schweitzer MH & Wittmeyer JL & Horner JR" box="[1088,1536,1801,1827]" journalOrPublisher="Science" pageId="12" pageNumber="13" pagination="1456 - 1460" part="308" refId="ref55976" refString="Schweitzer MH, Wittmeyer JL, Horner JR. 2005. Gender-specific reproductive tissue in ratites and Tyrannosaurus rex. Science 308 (5727): 1456 - 1460 DOI 10.1126 / science. 1112158." title="Gender-specific reproductive tissue in ratites and Tyrannosaurus rex" type="journal article" year="2005">
|
||
<emphasis box="[1088,1536,1801,1827]" italics="true" pageId="12" pageNumber="13">Schweitzer, Wittmeyer & Horner, 2005</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[498,763,1841,1867]" journalOrPublisher="Science Advances" pageId="12" pageNumber="13" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[498,763,1841,1867]" italics="true" pageId="12" pageNumber="13">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
; this assessment has been challenged by
|
||
<emphasis box="[1247,1497,1841,1867]" italics="true" pageId="12" pageNumber="13">
|
||
O’
|
||
<bibRefCitation author="O ' Connor J & Erickson GM & Norell M & Bailleul AM & Hu H & Zhou Z." box="[1273,1497,1841,1867]" journalOrPublisher="Nature Communications" pageId="12" pageNumber="13" pagination="443" part="9" refId="ref55359" refString="O ' Connor J, Erickson GM, Norell M, Bailleul AM, Hu H, Zhou Z. 2018. Medullary bone in an early Cretaceous enantiornithine bird and discussion regarding its identification in fossils. Nature Communications 9 (1): 443 DOI 10.1038 / s 41467 - 018 - 07621 - z." title="Medullary bone in an early Cretaceous enantiornithine bird and discussion regarding its identification in fossils" type="journal article" year="2018">Connor et al., 2018</bibRefCitation>
|
||
</emphasis>
|
||
), were recovered as a subadult and a young adult, respectively (
|
||
<figureCitation box="[1225,1290,1880,1907]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="12" pageNumber="13">Fig. 2</figureCitation>
|
||
). Sexual dimorphism was presumed here to be expressed in one of two ways: either the ontogram would split into separate male and female branches, or sexually diagnostic characters might be optimized as individual variation.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824976" ID-Zenodo-Dep="5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="13" pageNumber="14" startId="13.[524,589,1473,1495]" subCaptionStartIDs="13.[524,556,1737,1759]" subCaptionStarts="Fig. 12" targetBox="[475,1542,220,1456]" targetPageId="13">
|
||
<paragraph pageId="13" pageNumber="14">
|
||
<emphasis bold="true" pageId="13" pageNumber="14">
|
||
Figure 2 Ontogram of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[770,960,1474,1495]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[770,960,1474,1495]" italics="true" pageId="13" pageNumber="14">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages.
|
||
</emphasis>
|
||
Arrowhead points to the most mature specimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1035,1090,1679,1700]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1035,1090,1679,1700]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is metamorphic (sensu
|
||
<bibRefCitation author="Rose CS & Reiss JO" box="[1323,1502,1678,1700]" editor="Hanken H & Hall BK" journalOrPublisher="Chicago: University of Chicago Press" pageId="13" pageNumber="14" refId="ref55796" refString="Rose CS, Reiss JO. 1993. Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms. In: Hanken H, Hall BK, eds. The Skull: Development. Vol. 1. Chicago: University of Chicago Press." title="Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms" type="book" volumeTitle="The Skull: Development. Vol. 1" year="1993">
|
||
<emphasis box="[1323,1502,1678,1700]" italics="true" pageId="13" pageNumber="14">Rose & Reiss, 1993</emphasis>
|
||
</bibRefCitation>
|
||
). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1272,1326,1737,1758]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1272,1326,1737,1758]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale;
|
||
<materialsCitation box="[524,715,1824,1846]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="13" pageNumber="14" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFCAFFA6FF93FF7032F6FEDA" httpUri="http://table.plazi.org/id/DF5C84E2FFCAFFA6FF93FF7032F6FEDA" pageId="14" pageNumber="15" startId="14.[117,172,248,269]" targetBox="[108,1525,378,705]" targetIsTable="true" targetPageId="14">
|
||
<paragraph blockId="14.[117,1516,248,358]" pageId="14" pageNumber="15">
|
||
<emphasis bold="true" pageId="14" pageNumber="15">
|
||
Table 2 An assessment of evidence for sexual dimorphism in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[728,914,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[728,914,249,270]" italics="true" pageId="14" pageNumber="15">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
based on character states that are unambiguously optimized as individual variation.
|
||
</emphasis>
|
||
The female specimen
|
||
<materialsCitation box="[558,663,277,299]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
shares homologous character transformations with five other specimens, but these shared changes are not common to all of the specimens, which would be expected if they represented diagnostic character states unique to one sex. Based on this lack of pattern it is inferred that character states unique to females are absent and, by extension, the skeleton of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1211,1265,337,358]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1211,1265,337,358]" italics="true" pageId="14" pageNumber="15">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is not sexually dimorphic.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="14" pageNumber="15">
|
||
<table box="[108,1525,378,705]" gridcols="6" gridrows="9" pageId="14" pageNumber="15">
|
||
<tr box="[108,1525,378,400]" gridrow="0" pageId="14" pageNumber="15">
|
||
<th box="[108,576,378,400]" gridcol="0" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[108,218,378,400]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<th box="[621,749,378,400]" gridcol="1" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[621,749,378,400]" collectionCode="CMNH" pageId="14" pageNumber="15" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
</th>
|
||
<th box="[793,939,378,400]" gridcol="2" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[793,939,378,400]" collectionCode="LACM" pageId="14" pageNumber="15" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</th>
|
||
<th box="[983,1082,378,400]" gridcol="3" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[983,1082,378,400]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<th box="[1126,1283,378,400]" gridcol="4" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[1126,1283,378,400]" collectionCode="BMRP" pageId="14" pageNumber="15" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<th box="[1327,1525,378,400]" gridcol="5" gridrow="0" pageId="14" pageNumber="15">
|
||
<materialsCitation box="[1327,1525,378,399]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="14" pageNumber="15" specimenCode="AMNH FARB 5117">
|
||
<emphasis bold="true" box="[1327,1525,378,399]" pageId="14" pageNumber="15">AMNH FARB 5117</emphasis>
|
||
</materialsCitation>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1525,419,441]" gridrow="1" pageId="14" pageNumber="15" rowspan-1="1" rowspan-3="1" rowspan-4="1" rowspan-5="1">
|
||
<th box="[108,576,419,441]" gridcol="0" gridrow="1" pageId="14" pageNumber="15">330. Maxilla, subnrl frmn, dpth, 0 => 1</th>
|
||
<td box="[793,939,419,441]" gridcol="2" gridrow="1" pageId="14" pageNumber="15">330, 0=>1</td>
|
||
</tr>
|
||
<tr box="[108,1525,457,479]" gridrow="2" pageId="14" pageNumber="15" rowspan-1="1" rowspan-2="1" rowspan-4="1" rowspan-5="1">
|
||
<th box="[108,576,457,479]" gridcol="0" gridrow="2" pageId="14" pageNumber="15">419. Maxilla, js for jgl, rstrl extnt, 0 => 1</th>
|
||
<td box="[983,1082,457,479]" gridcol="3" gridrow="2" pageId="14" pageNumber="15">419, 0=>1</td>
|
||
</tr>
|
||
<tr box="[108,1525,495,517]" gridrow="3" pageId="14" pageNumber="15" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-5="1">
|
||
<th box="[108,576,495,517]" gridcol="0" gridrow="3" pageId="14" pageNumber="15">449. Lacrimal, drsl rms, sbctns srfc, hght, 0 => 1</th>
|
||
<td box="[1126,1283,495,517]" gridcol="4" gridrow="3" pageId="14" pageNumber="15">449, 0=>1</td>
|
||
</tr>
|
||
<tr box="[108,1525,532,554]" gridrow="4" pageId="14" pageNumber="15" rowspan-1="1" rowspan-3="1" rowspan-4="1" rowspan-5="1">
|
||
<th box="[108,576,532,554]" gridcol="0" gridrow="4" pageId="14" pageNumber="15">618. Postorbital, drsl mrgn, ornnttn, 1 => 2</th>
|
||
<td box="[793,939,532,554]" gridcol="2" gridrow="4" pageId="14" pageNumber="15">618, 0=>1</td>
|
||
</tr>
|
||
<tr box="[108,1525,570,592]" gridrow="5" pageId="14" pageNumber="15" rowspan-1="1" rowspan-2="1" rowspan-4="1" rowspan-5="1">
|
||
<th box="[108,576,570,592]" gridcol="0" gridrow="5" pageId="14" pageNumber="15">728. Quadratojugal, sq pr, rstrl srfc, 1 => 2</th>
|
||
<td box="[983,1082,570,592]" gridcol="3" gridrow="5" pageId="14" pageNumber="15">728, 1=>2</td>
|
||
</tr>
|
||
<tr box="[108,1525,607,629]" gridrow="6" pageId="14" pageNumber="15" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1">
|
||
<th box="[108,576,607,629]" gridcol="0" gridrow="6" pageId="14" pageNumber="15">963. Parietal, sggtl crst, cdl end, extnt, 0 ==> 1</th>
|
||
<td box="[621,749,607,629]" gridcol="1" gridrow="6" pageId="14" pageNumber="15">963, 0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[108,1525,645,667]" gridrow="7" pageId="14" pageNumber="15" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1">
|
||
<th box="[108,576,645,667]" gridcol="0" gridrow="7" pageId="14" pageNumber="15">973. Parietal, prtsoc pllr, prsnc, 1 => 2</th>
|
||
<td box="[1327,1525,645,667]" gridcol="5" gridrow="7" pageId="14" pageNumber="15">973, 1=>2</td>
|
||
</tr>
|
||
<tr box="[108,1525,683,705]" gridrow="8" pageId="14" pageNumber="15" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1">
|
||
<th box="[108,576,683,705]" gridcol="0" gridrow="8" pageId="14" pageNumber="15">1022. Basioccipital, occ con, cdl mrgn, 0 => 1</th>
|
||
<td box="[1327,1525,683,705]" gridcol="5" gridrow="8" pageId="14" pageNumber="15">1022, 0=>1</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="14.[498,1542,780,1444]" pageId="14" pageNumber="15">
|
||
In the first instance, a split of the ontogram into separate branches is not seen. In the second, the individual variation of
|
||
<materialsCitation box="[899,1027,939,966]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
was compared with that of other specimens. Sexually informative variation, in this case female variation, was assumed to be expressed as multiple homologous character changes seen in
|
||
<materialsCitation box="[1099,1229,1019,1046]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
and a repeated set of specimens (i.e., the identical set of specimens for each shared homologous character change). The comparison found eight homologous character changes shared between
|
||
<materialsCitation box="[498,626,1138,1165]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
and five other specimens; however, the character changes are not shared with a uniform set of specimens (
|
||
<tableCitation box="[814,902,1179,1205]" captionStart="Table 2" captionStartId="14.[117,172,248,269]" captionTargetBox="[108,1525,378,705]" captionText="Table 2 An assessment of evidence for sexual dimorphism in Tyrannosaurus rex based on character states that are unambiguously optimized as individual variation. The female specimen MOR 1125 shares homologous character transformations with five other specimens, but these shared changes are not common to all of the specimens, which would be expected if they represented diagnostic character states unique to one sex. Based on this lack of pattern it is inferred that character states unique to females are absent and, by extension, the skeleton of T. rex is not sexually dimorphic." httpUri="http://table.plazi.org/id/DF5C84E2FFCAFFA6FF93FF7032F6FEDA" pageId="14" pageNumber="15" tableUuid="DF5C84E2FFCAFFA6FF93FF7032F6FEDA">Table 2</tableCitation>
|
||
). Therefore, there is no skeletodental evidence for sexual dimorphism in the data set (i.e., males and females are skeletally identical aside from the presence of medullary bone); otherwise, the ontogram would have divided into separate male and female branches or homologous character changes would have singled out
|
||
<materialsCitation box="[546,676,1338,1365]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
and an associated set of specimens. This test is not dependent upon
|
||
<materialsCitation box="[498,626,1377,1404]" collectionCode="MOR" pageId="14" pageNumber="15" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
being a female (
|
||
<bibRefCitation author="O ' Connor J & Erickson GM & Norell M & Bailleul AM & Hu H & Zhou Z." box="[816,1060,1378,1404]" journalOrPublisher="Nature Communications" pageId="14" pageNumber="15" pagination="443" part="9" refId="ref55359" refString="O ' Connor J, Erickson GM, Norell M, Bailleul AM, Hu H, Zhou Z. 2018. Medullary bone in an early Cretaceous enantiornithine bird and discussion regarding its identification in fossils. Nature Communications 9 (1): 443 DOI 10.1038 / s 41467 - 018 - 07621 - z." title="Medullary bone in an early Cretaceous enantiornithine bird and discussion regarding its identification in fossils" type="journal article" year="2018">
|
||
<emphasis box="[816,1060,1378,1404]" italics="true" pageId="14" pageNumber="15">O’ Connor et al., 2018</emphasis>
|
||
</bibRefCitation>
|
||
); if in actuality it is a male, then the set of shared characters would be evidence of that sex.
|
||
</paragraph>
|
||
<paragraph blockId="14.[498,1542,1489,1920]" box="[498,804,1489,1519]" pageId="14" pageNumber="15">
|
||
<heading bold="true" box="[498,804,1489,1519]" fontSize="12" level="2" pageId="14" pageNumber="15" reason="0">
|
||
<emphasis bold="true" box="[498,804,1489,1519]" pageId="14" pageNumber="15">Wildcard specimens</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="14.[498,1542,1489,1920]" pageId="14" pageNumber="15">
|
||
Thirteen wildcard specimens were identified; in each case, the analyses that included them resulted in multiple ontograms that collapsed into one or several polytomies; the regions of collapse are enclosed by ellipses in
|
||
<figureCitation box="[913,996,1614,1640]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="14" pageNumber="15">Fig. 1B</figureCitation>
|
||
. Upon comparison of the polytomies, the wildcard specimens could be divided into two groups, where resolution was lost either rootward or distalward relative to
|
||
<materialsCitation box="[903,1088,1694,1721]" collectionCode="BMRP" pageId="14" pageNumber="15" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
, the most complete specimen in the dataset; that is, that specimen was not part of either polytomy. The partial adult specimen,
|
||
<materialsCitation box="[498,610,1774,1800]" collectionCode="CM" pageId="14" pageNumber="15" specimenCode="CM 1400">CM 1400</materialsCitation>
|
||
, is shown in a polytomy with two other specimens at the end of a branch (
|
||
<figureCitation box="[508,591,1813,1840]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="14" pageNumber="15">Fig. 1B</figureCitation>
|
||
); however, structure was lost elsewhere in the ontogram when this specimen was included in the analysis, and so it was excluded from the backbone topology (
|
||
<figureCitation box="[1450,1542,1853,1879]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="14" pageNumber="15">Figs. 1B</figureCitation>
|
||
and
|
||
<figureCitation box="[551,565,1893,1919]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="14" pageNumber="15">2</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,234,540]" pageId="15" pageNumber="16">
|
||
Despite the fact that the wildcards are coded for less than 7% of the characters, some occupy a relatively precise location of the ontogram (e.g.,
|
||
<materialsCitation box="[1231,1341,274,300]" collectionCode="CM" pageId="15" pageNumber="16" specimenCode="CM 1400">CM 1400</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1352,1462,274,300]" collectionCode="CM" pageId="15" pageNumber="16" specimenCode="CM 9401">CM 9401</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 35.1">DDM 35.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[562,694,314,340]" collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 3044">MOR 3044</materialsCitation>
|
||
,
|
||
<materialsCitation box="[707,839,314,341]" collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 2925">MOR 2925</materialsCitation>
|
||
,
|
||
<materialsCitation box="[852,1000,314,340]" collectionCode="RSM" pageId="15" pageNumber="16" specimenCode="RSM 2347.1">RSM 2347.1</materialsCitation>
|
||
), whereas others collapse entire regions (e.g.,
|
||
<materialsCitation box="[560,758,354,380]" collectionCode="BMRP" pageId="15" pageNumber="16" specimenCode="BMRP 2001.4.70">BMRP 2001.4.70</materialsCitation>
|
||
, DDM 1536.1,
|
||
<materialsCitation box="[935,1104,354,381]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 1562.14">DDM 1562.14</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1115,1268,354,380]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 1863.1">DDM 1863.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1280,1465,354,380]" collectionCode="FMNH" pageId="15" pageNumber="16" specimenCode="FMNH PR2411">FMNH PR2411</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 002">MOR 002</materialsCitation>
|
||
,
|
||
<materialsCitation box="[555,732,394,421]" collectionCode="TMM" pageId="15" pageNumber="16" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
|
||
). Of these specimens, many are single bones (
|
||
<materialsCitation box="[1274,1476,394,420]" collectionCode="BMRP" pageId="15" pageNumber="16" specimenCode="BMRP 2001.4.70">BMRP 2001.4.70</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,626,433,460]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 35.1">DDM 35.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[639,794,433,460]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 1536.8">DDM 1536.8</materialsCitation>
|
||
,
|
||
<materialsCitation box="[808,994,434,460]" collectionCode="FMNH" pageId="15" pageNumber="16" specimenCode="FMNH PR2411">FMNH PR2411</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1007,1139,433,460]" collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 2925">MOR 2925</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1152,1284,434,460]" collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 3044">MOR 3044</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1297,1442,434,460]" collectionCode="RSM" pageId="15" pageNumber="16" specimenCode="RSM 2347.1">RSM 2347.1</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="TMM" pageId="15" pageNumber="16" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
|
||
) or teeth (
|
||
<materialsCitation box="[715,885,473,500]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 1562.14">DDM 1562.14</materialsCitation>
|
||
,
|
||
<materialsCitation box="[896,1051,473,500]" collectionCode="DDM" pageId="15" pageNumber="16" specimenCode="DDM 1863.1">DDM 1863.1</materialsCitation>
|
||
), whereas only two are a partial skull (
|
||
<materialsCitation collectionCode="CM" pageId="15" pageNumber="16" specimenCode="CM 1400">CM 1400</materialsCitation>
|
||
) or skull and skeleton (
|
||
<materialsCitation box="[835,959,513,539]" collectionCode="MOR" pageId="15" pageNumber="16" specimenCode="MOR 002">MOR 002</materialsCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,576,1089]" box="[498,1156,576,606]" pageId="15" pageNumber="16">
|
||
<heading bold="true" box="[498,1156,576,606]" fontSize="12" level="2" pageId="15" pageNumber="16" reason="0">
|
||
<emphasis bold="true" box="[498,1156,576,606]" pageId="15" pageNumber="16">Overview of frequencies of growth changes</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,576,1089]" box="[498,1004,621,649]" pageId="15" pageNumber="16">
|
||
<heading bold="true" box="[498,1004,621,649]" fontSize="11" level="3" pageId="15" pageNumber="16" reason="6">
|
||
<emphasis bold="true" box="[498,1004,621,649]" italics="true" pageId="15" pageNumber="16">Influence of specimen completeness</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,576,1089]" pageId="15" pageNumber="16">
|
||
The effect of specimen completeness upon the number of synontomorphies at each node was quantified through a Spearman rank correlation test. Examination of the raw data as histograms with a fitted normal curve showed they are positively skewed, and a Shapiro
|
||
<emphasis box="[588,603,785,810]" italics="true" pageId="15" pageNumber="16">–</emphasis>
|
||
Wilk test of normality found that they were not normally distributed (completeness,
|
||
<emphasis box="[677,691,825,850]" italics="true" pageId="15" pageNumber="16">p</emphasis>
|
||
= 0.001; synontomorphies,
|
||
<emphasis box="[1012,1026,825,850]" italics="true" pageId="15" pageNumber="16">p</emphasis>
|
||
= 0.000) and so the data were normalized by converting them to ranks (
|
||
<tableCitation box="[816,903,864,890]" captionStart="Table 3" captionStartId="16.[117,172,248,269]" captionTargetBox="[108,1508,319,1569]" captionText="Table 3 Comparison of the completeness of Tyrannosaurus rex specimens included in the backbone ontogram and the number of synontomorphies at each node. Raw data are in normal typeface, ranks used in the correlation test are in boldface." httpUri="http://table.plazi.org/id/DF5C84E2FFD4FFB8FF93FF7033CFFE97" pageId="15" pageNumber="16" tableUuid="DF5C84E2FFD4FFB8FF93FF7033CFFE97">Table 3</tableCitation>
|
||
;
|
||
<figureCitation box="[918,984,864,890]" captionStart="Figure 3" captionStartId="17.[524,589,657,679]" captionTargetBox="[473,1323,220,638]" captionTargetId="figure-243@17.[497,1066,233,631]" captionTargetPageId="17" captionText="Figure 3 Scatterplot showing the noncongruence in Tyrannosaurus rex between the completeness of specimens (i.e., number of characters scored) and the number of synontomorphies at each corresponding node. Per cent completeness (decreasing away from the origin) and the number of synontomorphies supporting the corresponding node (decreasing away from the origin) have been converted to ranks. A Spearman correlation test on these data results in a nonsignificant correlation coefficient; ergo, the number of synontomorphies at an internode is not an artifact of specimen completeness. Full-size DOI: 10.7717/peerj.9192/fig-3" figureDoi="http://doi.org/10.5281/zenodo.5824978" httpUri="https://zenodo.org/record/5824978/files/figure.png" pageId="15" pageNumber="16">Fig. 3</figureCitation>
|
||
). A Shapiro-Wilk test of normality found that the ranked completeness and ranked number of synontomorphies were normally distributed (
|
||
<emphasis box="[643,657,944,969]" italics="true" pageId="15" pageNumber="16">p</emphasis>
|
||
= 0.217, 0.134, respectively). A Spearman rank correlation test resulted in a nonsignificant (
|
||
<emphasis box="[683,697,984,1009]" italics="true" pageId="15" pageNumber="16">p</emphasis>
|
||
= 0.423) correlation coefficient (
|
||
<emphasis box="[1082,1093,984,1009]" italics="true" pageId="15" pageNumber="16">
|
||
<emphasis box="[1093,1103,996,1012]" italics="true" pageId="15" pageNumber="16">
|
||
<subScript attach="left" box="[1093,1103,996,1012]" fontSize="7" pageId="15" pageNumber="16">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.149), indicating that the peaks of synontomorphies at each node are not dependent upon the completeness of the specimens that extend from each branch.
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,1128,1916]" box="[498,816,1128,1156]" pageId="15" pageNumber="16">
|
||
<heading bold="true" box="[498,816,1128,1156]" fontSize="11" level="3" pageId="15" pageNumber="16" reason="6">
|
||
<emphasis bold="true" box="[498,816,1128,1156]" italics="true" pageId="15" pageNumber="16">Synontomorphy trends</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,1128,1916]" pageId="15" pageNumber="16">
|
||
The overall distribution of growth changes is unimodal (
|
||
<figureCitation box="[1168,1233,1172,1198]" captionStart="Figure 4" captionStartId="17.[524,589,1366,1388]" captionTargetBox="[500,1530,910,1333]" captionTargetId="figure-134@17.[497,1129,908,1339]" captionTargetPageId="17" captionText="Figure 4 Frequency distribution of unambiguously optimized synontomorphies during the growth of Tyrannosaurus rex. Growth stages (corresponding to the numbered nodes of the ontogram in Fig. 2) are along the x-axis and the number of changes are along the y-axis. The greatest number of changes are seen in the transition from large juvenile to subadult, or, from growth stage 5–6; the high concentration of change between these growth categories is evidence that T. rex ontogeny is metamorphic (sensu Rose & Reiss, 1993). Full-size DOI: 10.7717/peerj.9192/fig-4" figureDoi="http://doi.org/10.5281/zenodo.5824980" httpUri="https://zenodo.org/record/5824980/files/figure.png" pageId="15" pageNumber="16">Fig. 4</figureCitation>
|
||
): the highest number occurs early, peaking at the subadult growth category, which then precipitously drops, aside from several low peaks in the adult growth stages. In total, there are five peaks, and four of them mark the onset of growth categories. The peak at growth stage 6 marks the beginning of the subadult growth category, where an increase in the height of the skull frame and inflation of the bones that enclose the antorbital sinus are seen, as well as changes to the pectoral girdle and limb, and pes. The peak at growth stage 8 marks the onset of the young adult growth category. The peak at growth stage 13 marks the onset of adulthood, where the External Fundamental System (EFS; a narrow band of lines of arrested growth that indicate near cessation of appositional growth) is first seen as well as changes to the antorbital sinus system and the origin of the adductor musculature. The peak at growth stage 19 marks extensive changes to the skull and postcranial skeleton, but by itself does not define a new growth category. Finally, the peak of changes at growth stage 21 correspond to the extensive changes in the skeleton of
|
||
<materialsCitation box="[1250,1436,1690,1717]" collectionCode="FMNH" pageId="15" pageNumber="16" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
, the most mature and second-best sampled specimen in the data set.
|
||
</paragraph>
|
||
<paragraph blockId="15.[498,1542,1128,1916]" lastBlockId="16.[498,1541,1639,1904]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
|
||
Given their abundance in the character matrix, nonphylogenetic synontomorphies are more numerous at each node than phylogenetic synontomorphies, but they broadly follow the same pattern of peaks and valleys (
|
||
<figureCitation box="[1023,1086,1849,1876]" captionStart="Figure 5" captionStartId="18.[524,589,692,714]" captionTargetBox="[498,1541,234,679]" captionTargetId="figure-212@18.[497,1128,233,666]" captionTargetPageId="18" captionText="Figure 5 Comparison of the frequency distributions of phylogenetic and nonphylogenetic synontomorphies in the ontogeny of Tyrannosaurus rex. Growth stage is along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and number of synontomorphies is along the y-axis. Phylogenetic characters are in solid bars; nonphylogenetic characters are in hollow bars. The frequency distributions of both sets of data follow the same general pattern, aside from the flatter distribution of the phylogenetic synontomorphies relative to the nonphylogenetic synontomorphies and the reversed pattern seen at growth stages 7 and 8. Both types of changes occur throughout the lifespan of T. rex, indicating that ontogeny is not strictly congruent with phylogeny. Full-size DOI: 10.7717/peerj.9192/fig-5" figureDoi="http://doi.org/10.5281/zenodo.5824982" httpUri="https://zenodo.org/record/5824982/files/figure.png" pageId="15" pageNumber="16">Fig. 5</figureCitation>
|
||
), aside from growth stages 6, 8, 10, and 21, where phylogenetic changes tend to decrease in frequency whereas nonphylogenetic changes increase. This difference in frequency distribution shows that the number of phylogenetic changes is not controlled by the number of characters scored. Phylogenetic changes are frequent (i.e., more than five) early in growth (stages 2
|
||
<emphasis box="[1290,1305,1719,1744]" italics="true" pageId="16" pageNumber="17">–</emphasis>
|
||
9), whereas they are less common among adults (stages 10, 12, 13, 18
|
||
<emphasis box="[1074,1089,1759,1784]" italics="true" pageId="16" pageNumber="17">–</emphasis>
|
||
21); if ontogeny is congruent with phylogeny then this gross pattern should be expected since fewer phylogenetic synontomorphies should occur among adults, which presumably would only express characters at the level of species.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFD4FFB8FF93FF7033CFFE97" httpUri="http://table.plazi.org/id/DF5C84E2FFD4FFB8FF93FF7033CFFE97" pageId="16" pageNumber="17" startId="16.[117,172,248,269]" targetBox="[108,1508,319,1569]" targetIsTable="true" targetPageId="16">
|
||
<paragraph blockId="16.[117,1515,248,299]" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" pageId="16" pageNumber="17">
|
||
Table 3 Comparison of the completeness of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[598,790,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[598,790,249,270]" italics="true" pageId="16" pageNumber="17">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
specimens included in the backbone ontogram and the number of synontomorphies at each node.
|
||
</emphasis>
|
||
Raw data are in normal typeface, ranks used in the correlation test are in boldface.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="16" pageNumber="17">
|
||
<table box="[108,1508,319,1569]" gridcols="7" gridrows="32" pageId="16" pageNumber="17">
|
||
<tr box="[108,1508,319,400]" gridrow="0" pageId="16" pageNumber="17">
|
||
<th box="[108,301,319,400]" gridcol="0" gridrow="0" pageId="16" pageNumber="17">Specimen</th>
|
||
<th box="[329,461,319,400]" gridcol="1" gridrow="0" pageId="16" pageNumber="17">Specimen completeness</th>
|
||
<th box="[512,697,319,400]" gridcol="2" gridrow="0" pageId="16" pageNumber="17">Specimen completeness rank</th>
|
||
<th box="[733,865,319,400]" gridcol="3" gridrow="0" pageId="16" pageNumber="17">Specimen completeness midranks</th>
|
||
<th box="[887,1083,319,400]" gridcol="4" gridrow="0" pageId="16" pageNumber="17"># Synontomorphies at node</th>
|
||
<th box="[1101,1290,319,400]" gridcol="5" gridrow="0" pageId="16" pageNumber="17">#Synontomorphies rank</th>
|
||
<th box="[1313,1508,319,400]" gridcol="6" gridrow="0" pageId="16" pageNumber="17">
|
||
# Synontomorphies
|
||
<emphasis bold="true" box="[1313,1408,349,371]" pageId="16" pageNumber="17">midranks</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1508,419,441]" gridrow="1" pageId="16" pageNumber="17">
|
||
<th box="[108,301,419,441]" gridcol="0" gridrow="1" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,257,420,441]" collectionCode="BMRP" pageId="16" pageNumber="17" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,419,441]" gridcol="1" gridrow="1" pageId="16" pageNumber="17">86.7</td>
|
||
<td box="[512,697,419,441]" gridcol="2" gridrow="1" pageId="16" pageNumber="17">1</td>
|
||
<td box="[733,865,419,441]" gridcol="3" gridrow="1" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,419,440]" pageId="16" pageNumber="17">1</emphasis>
|
||
</td>
|
||
<td box="[887,1083,419,441]" gridcol="4" gridrow="1" pageId="16" pageNumber="17">51</td>
|
||
<td box="[1101,1290,419,441]" gridcol="5" gridrow="1" pageId="16" pageNumber="17">3</td>
|
||
<td box="[1313,1508,419,441]" gridcol="6" gridrow="1" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1325,419,440]" pageId="16" pageNumber="17">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,457,479]" gridrow="2" pageId="16" pageNumber="17">
|
||
<th box="[108,301,457,479]" gridcol="0" gridrow="2" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,258,457,479]" collectionCode="FMNH" pageId="16" pageNumber="17" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,457,479]" gridcol="1" gridrow="2" pageId="16" pageNumber="17">84.3</td>
|
||
<td box="[512,697,457,479]" gridcol="2" gridrow="2" pageId="16" pageNumber="17">2</td>
|
||
<td box="[733,865,457,479]" gridcol="3" gridrow="2" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,457,478]" pageId="16" pageNumber="17">2</emphasis>
|
||
</td>
|
||
<td box="[887,1083,457,479]" gridcol="4" gridrow="2" pageId="16" pageNumber="17">7</td>
|
||
<td box="[1101,1290,457,479]" gridcol="5" gridrow="2" pageId="16" pageNumber="17">18</td>
|
||
<td box="[1313,1508,457,479]" gridcol="6" gridrow="2" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,457,478]" pageId="16" pageNumber="17">19</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,494,516]" gridrow="3" pageId="16" pageNumber="17">
|
||
<th box="[108,301,494,516]" gridcol="0" gridrow="3" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,214,494,516]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,494,516]" gridcol="1" gridrow="3" pageId="16" pageNumber="17">57.8</td>
|
||
<td box="[512,697,494,516]" gridcol="2" gridrow="3" pageId="16" pageNumber="17">3</td>
|
||
<td box="[733,865,494,516]" gridcol="3" gridrow="3" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,494,515]" pageId="16" pageNumber="17">3</emphasis>
|
||
</td>
|
||
<td box="[887,1083,494,516]" gridcol="4" gridrow="3" pageId="16" pageNumber="17">31</td>
|
||
<td box="[1101,1290,494,516]" gridcol="5" gridrow="3" pageId="16" pageNumber="17">4</td>
|
||
<td box="[1313,1508,494,516]" gridcol="6" gridrow="3" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1325,494,515]" pageId="16" pageNumber="17">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,532,554]" gridrow="4" pageId="16" pageNumber="17">
|
||
<th box="[108,301,532,554]" gridcol="0" gridrow="4" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,202,532,553]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,532,554]" gridcol="1" gridrow="4" pageId="16" pageNumber="17">57.6</td>
|
||
<td box="[512,697,532,554]" gridcol="2" gridrow="4" pageId="16" pageNumber="17">4</td>
|
||
<td box="[733,865,532,554]" gridcol="3" gridrow="4" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,532,553]" pageId="16" pageNumber="17">4</emphasis>
|
||
</td>
|
||
<td box="[887,1083,532,554]" gridcol="4" gridrow="4" pageId="16" pageNumber="17">2</td>
|
||
<td box="[1101,1290,532,554]" gridcol="5" gridrow="4" pageId="16" pageNumber="17">25</td>
|
||
<td box="[1313,1508,532,554]" gridcol="6" gridrow="4" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,532,553]" pageId="16" pageNumber="17">26.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,570,592]" gridrow="5" pageId="16" pageNumber="17">
|
||
<th box="[108,301,570,592]" gridcol="0" gridrow="5" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,250,570,592]" collectionCode="LACM" pageId="16" pageNumber="17" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,570,592]" gridcol="1" gridrow="5" pageId="16" pageNumber="17">55.9</td>
|
||
<td box="[512,697,570,592]" gridcol="2" gridrow="5" pageId="16" pageNumber="17">5</td>
|
||
<td box="[733,865,570,592]" gridcol="3" gridrow="5" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,570,591]" pageId="16" pageNumber="17">5</emphasis>
|
||
</td>
|
||
<td box="[887,1083,570,592]" gridcol="4" gridrow="5" pageId="16" pageNumber="17">12</td>
|
||
<td box="[1101,1290,570,592]" gridcol="5" gridrow="5" pageId="16" pageNumber="17">12</td>
|
||
<td box="[1313,1508,570,592]" gridcol="6" gridrow="5" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,570,591]" pageId="16" pageNumber="17">12</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,607,629]" gridrow="6" pageId="16" pageNumber="17">
|
||
<th box="[108,301,607,629]" gridcol="0" gridrow="6" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,247,608,629]" collectionCode="UWBM" pageId="16" pageNumber="17" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,607,629]" gridcol="1" gridrow="6" pageId="16" pageNumber="17">50.7</td>
|
||
<td box="[512,697,607,629]" gridcol="2" gridrow="6" pageId="16" pageNumber="17">6</td>
|
||
<td box="[733,865,607,629]" gridcol="3" gridrow="6" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,607,628]" pageId="16" pageNumber="17">6</emphasis>
|
||
</td>
|
||
<td box="[887,1083,607,629]" gridcol="4" gridrow="6" pageId="16" pageNumber="17">9</td>
|
||
<td box="[1101,1290,607,629]" gridcol="5" gridrow="6" pageId="16" pageNumber="17">15</td>
|
||
<td box="[1313,1508,607,629]" gridcol="6" gridrow="6" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,607,628]" pageId="16" pageNumber="17">16</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,645,667]" gridrow="7" pageId="16" pageNumber="17">
|
||
<th box="[108,301,645,667]" gridcol="0" gridrow="7" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,202,645,667]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,645,667]" gridcol="1" gridrow="7" pageId="16" pageNumber="17">47.3</td>
|
||
<td box="[512,697,645,667]" gridcol="2" gridrow="7" pageId="16" pageNumber="17">7</td>
|
||
<td box="[733,865,645,667]" gridcol="3" gridrow="7" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,645,666]" pageId="16" pageNumber="17">7</emphasis>
|
||
</td>
|
||
<td box="[887,1083,645,667]" gridcol="4" gridrow="7" pageId="16" pageNumber="17">2</td>
|
||
<td box="[1101,1290,645,667]" gridcol="5" gridrow="7" pageId="16" pageNumber="17">26</td>
|
||
<td box="[1313,1508,645,667]" gridcol="6" gridrow="7" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,645,666]" pageId="16" pageNumber="17">26.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,683,705]" gridrow="8" pageId="16" pageNumber="17">
|
||
<th box="[108,301,683,705]" gridcol="0" gridrow="8" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,202,683,704]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,683,705]" gridcol="1" gridrow="8" pageId="16" pageNumber="17">42.4</td>
|
||
<td box="[512,697,683,705]" gridcol="2" gridrow="8" pageId="16" pageNumber="17">8</td>
|
||
<td box="[733,865,683,705]" gridcol="3" gridrow="8" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,683,704]" pageId="16" pageNumber="17">8</emphasis>
|
||
</td>
|
||
<td box="[887,1083,683,705]" gridcol="4" gridrow="8" pageId="16" pageNumber="17">15</td>
|
||
<td box="[1101,1290,683,705]" gridcol="5" gridrow="8" pageId="16" pageNumber="17">8</td>
|
||
<td box="[1313,1508,683,705]" gridcol="6" gridrow="8" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,683,704]" pageId="16" pageNumber="17">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,720,742]" gridrow="9" pageId="16" pageNumber="17">
|
||
<th box="[108,301,720,742]" gridcol="0" gridrow="9" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,224,720,742]" collectionCode="RSM" pageId="16" pageNumber="17" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,720,742]" gridcol="1" gridrow="9" pageId="16" pageNumber="17">40.5</td>
|
||
<td box="[512,697,720,742]" gridcol="2" gridrow="9" pageId="16" pageNumber="17">9</td>
|
||
<td box="[733,865,720,742]" gridcol="3" gridrow="9" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,745,720,741]" pageId="16" pageNumber="17">9</emphasis>
|
||
</td>
|
||
<td box="[887,1083,720,742]" gridcol="4" gridrow="9" pageId="16" pageNumber="17">20</td>
|
||
<td box="[1101,1290,720,742]" gridcol="5" gridrow="9" pageId="16" pageNumber="17">5</td>
|
||
<td box="[1313,1508,720,742]" gridcol="6" gridrow="9" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,720,741]" pageId="16" pageNumber="17">5.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,758,780]" gridrow="10" pageId="16" pageNumber="17">
|
||
<th box="[108,301,758,780]" gridcol="0" gridrow="10" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,299,758,780]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="16" pageNumber="17" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,758,780]" gridcol="1" gridrow="10" pageId="16" pageNumber="17">39.2</td>
|
||
<td box="[512,697,758,780]" gridcol="2" gridrow="10" pageId="16" pageNumber="17">10</td>
|
||
<td box="[733,865,758,780]" gridcol="3" gridrow="10" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,758,779]" pageId="16" pageNumber="17">10</emphasis>
|
||
</td>
|
||
<td box="[887,1083,758,780]" gridcol="4" gridrow="10" pageId="16" pageNumber="17">10</td>
|
||
<td box="[1101,1290,758,780]" gridcol="5" gridrow="10" pageId="16" pageNumber="17">13</td>
|
||
<td box="[1313,1508,758,780]" gridcol="6" gridrow="10" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,758,779]" pageId="16" pageNumber="17">13.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,795,817]" gridrow="11" pageId="16" pageNumber="17">
|
||
<th box="[108,301,795,817]" gridcol="0" gridrow="11" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,238,796,817]" collectionCode="LACM" pageId="16" pageNumber="17" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,795,817]" gridcol="1" gridrow="11" pageId="16" pageNumber="17">37.6</td>
|
||
<td box="[512,697,795,817]" gridcol="2" gridrow="11" pageId="16" pageNumber="17">11</td>
|
||
<td box="[733,865,795,817]" gridcol="3" gridrow="11" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,795,816]" pageId="16" pageNumber="17">11</emphasis>
|
||
</td>
|
||
<td box="[887,1083,795,817]" gridcol="4" gridrow="11" pageId="16" pageNumber="17">6</td>
|
||
<td box="[1101,1290,795,817]" gridcol="5" gridrow="11" pageId="16" pageNumber="17">21</td>
|
||
<td box="[1313,1508,795,817]" gridcol="6" gridrow="11" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,795,816]" pageId="16" pageNumber="17">21.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,833,855]" gridrow="12" pageId="16" pageNumber="17">
|
||
<th box="[108,301,833,855]" gridcol="0" gridrow="12" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,233,833,855]" collectionCode="CMNH" pageId="16" pageNumber="17" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,833,855]" gridcol="1" gridrow="12" pageId="16" pageNumber="17">28.2</td>
|
||
<td box="[512,697,833,855]" gridcol="2" gridrow="12" pageId="16" pageNumber="17">12</td>
|
||
<td box="[733,865,833,855]" gridcol="3" gridrow="12" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,833,854]" pageId="16" pageNumber="17">12</emphasis>
|
||
</td>
|
||
<td box="[887,1083,833,855]" gridcol="4" gridrow="12" pageId="16" pageNumber="17">3</td>
|
||
<td box="[1101,1290,833,855]" gridcol="5" gridrow="12" pageId="16" pageNumber="17">23</td>
|
||
<td box="[1313,1508,833,855]" gridcol="6" gridrow="12" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,833,854]" pageId="16" pageNumber="17">23.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,870,893]" gridrow="13" pageId="16" pageNumber="17">
|
||
<th box="[108,301,870,893]" gridcol="0" gridrow="13" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,198,871,892]" collectionCode="CM" pageId="16" pageNumber="17" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,870,893]" gridcol="1" gridrow="13" pageId="16" pageNumber="17">24.3</td>
|
||
<td box="[512,697,870,893]" gridcol="2" gridrow="13" pageId="16" pageNumber="17">13</td>
|
||
<td box="[733,865,870,893]" gridcol="3" gridrow="13" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,870,891]" pageId="16" pageNumber="17">13</emphasis>
|
||
</td>
|
||
<td box="[887,1083,870,893]" gridcol="4" gridrow="13" pageId="16" pageNumber="17">6</td>
|
||
<td box="[1101,1290,870,893]" gridcol="5" gridrow="13" pageId="16" pageNumber="17">22</td>
|
||
<td box="[1313,1508,870,893]" gridcol="6" gridrow="13" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,870,891]" pageId="16" pageNumber="17">21.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,908,930]" gridrow="14" pageId="16" pageNumber="17">
|
||
<th box="[108,301,908,930]" gridcol="0" gridrow="14" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,301,908,930]" collectionCode="TMP" pageId="16" pageNumber="17" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,908,930]" gridcol="1" gridrow="14" pageId="16" pageNumber="17">17.6</td>
|
||
<td box="[512,697,908,930]" gridcol="2" gridrow="14" pageId="16" pageNumber="17">14</td>
|
||
<td box="[733,865,908,930]" gridcol="3" gridrow="14" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,908,929]" pageId="16" pageNumber="17">14</emphasis>
|
||
</td>
|
||
<td box="[887,1083,908,930]" gridcol="4" gridrow="14" pageId="16" pageNumber="17">16</td>
|
||
<td box="[1101,1290,908,930]" gridcol="5" gridrow="14" pageId="16" pageNumber="17">7</td>
|
||
<td box="[1313,1508,908,930]" gridcol="6" gridrow="14" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1325,908,929]" pageId="16" pageNumber="17">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,946,968]" gridrow="15" pageId="16" pageNumber="17">
|
||
<th box="[108,301,946,968]" gridcol="0" gridrow="15" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,238,946,968]" collectionCode="LACM" pageId="16" pageNumber="17" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,946,968]" gridcol="1" gridrow="15" pageId="16" pageNumber="17">16.8</td>
|
||
<td box="[512,697,946,968]" gridcol="2" gridrow="15" pageId="16" pageNumber="17">15</td>
|
||
<td box="[733,865,946,968]" gridcol="3" gridrow="15" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,946,967]" pageId="16" pageNumber="17">15</emphasis>
|
||
</td>
|
||
<td box="[887,1083,946,968]" gridcol="4" gridrow="15" pageId="16" pageNumber="17">15</td>
|
||
<td box="[1101,1290,946,968]" gridcol="5" gridrow="15" pageId="16" pageNumber="17">9</td>
|
||
<td box="[1313,1508,946,968]" gridcol="6" gridrow="15" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,946,967]" pageId="16" pageNumber="17">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,983,1005]" gridrow="16" pageId="16" pageNumber="17">
|
||
<th box="[108,301,983,1005]" gridcol="0" gridrow="16" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,214,984,1005]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,983,1005]" gridcol="1" gridrow="16" pageId="16" pageNumber="17">16.7</td>
|
||
<td box="[512,697,983,1005]" gridcol="2" gridrow="16" pageId="16" pageNumber="17">16</td>
|
||
<td box="[733,865,983,1005]" gridcol="3" gridrow="16" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,983,1004]" pageId="16" pageNumber="17">16</emphasis>
|
||
</td>
|
||
<td box="[887,1083,983,1005]" gridcol="4" gridrow="16" pageId="16" pageNumber="17">7</td>
|
||
<td box="[1101,1290,983,1005]" gridcol="5" gridrow="16" pageId="16" pageNumber="17">19</td>
|
||
<td box="[1313,1508,983,1005]" gridcol="6" gridrow="16" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,983,1004]" pageId="16" pageNumber="17">19</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1021,1042]" gridrow="17" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1021,1042]" gridcol="0" gridrow="17" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,233,1021,1042]" collectionCode="SDSM" pageId="16" pageNumber="17" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1021,1042]" gridcol="1" gridrow="17" pageId="16" pageNumber="17">12.3</td>
|
||
<td box="[512,697,1021,1042]" gridcol="2" gridrow="17" pageId="16" pageNumber="17">17</td>
|
||
<td box="[733,865,1021,1042]" gridcol="3" gridrow="17" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1021,1042]" pageId="16" pageNumber="17">17</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1021,1042]" gridcol="4" gridrow="17" pageId="16" pageNumber="17">20</td>
|
||
<td box="[1101,1290,1021,1042]" gridcol="5" gridrow="17" pageId="16" pageNumber="17">6</td>
|
||
<td box="[1313,1508,1021,1042]" gridcol="6" gridrow="17" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,1021,1042]" pageId="16" pageNumber="17">5.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1058,1080]" gridrow="18" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1058,1080]" gridcol="0" gridrow="18" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,299,1058,1080]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="16" pageNumber="17" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1058,1080]" gridcol="1" gridrow="18" pageId="16" pageNumber="17">11.8</td>
|
||
<td box="[512,697,1058,1080]" gridcol="2" gridrow="18" pageId="16" pageNumber="17">18</td>
|
||
<td box="[733,865,1058,1080]" gridcol="3" gridrow="18" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1058,1079]" pageId="16" pageNumber="17">18</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1058,1080]" gridcol="4" gridrow="18" pageId="16" pageNumber="17">9</td>
|
||
<td box="[1101,1290,1058,1080]" gridcol="5" gridrow="18" pageId="16" pageNumber="17">16</td>
|
||
<td box="[1313,1508,1058,1080]" gridcol="6" gridrow="18" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,1058,1079]" pageId="16" pageNumber="17">16</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1096,1117]" gridrow="19" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1096,1117]" gridcol="0" gridrow="19" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,238,1096,1117]" collectionCode="LACM" pageId="16" pageNumber="17" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1096,1117]" gridcol="1" gridrow="19" pageId="16" pageNumber="17">10.4</td>
|
||
<td box="[512,697,1096,1117]" gridcol="2" gridrow="19" pageId="16" pageNumber="17">19</td>
|
||
<td box="[733,865,1096,1117]" gridcol="3" gridrow="19" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1096,1117]" pageId="16" pageNumber="17">19</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1096,1117]" gridcol="4" gridrow="19" pageId="16" pageNumber="17">0</td>
|
||
<td box="[1101,1290,1096,1117]" gridcol="5" gridrow="19" pageId="16" pageNumber="17">30</td>
|
||
<td box="[1313,1508,1096,1117]" gridcol="6" gridrow="19" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1096,1117]" pageId="16" pageNumber="17">30.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1134,1156]" gridrow="20" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1134,1156]" gridcol="0" gridrow="20" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,228,1134,1155]" collectionCode="UWGM" pageId="16" pageNumber="17" specimenCode="UWGM 181">UWGM 181</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1134,1156]" gridcol="1" gridrow="20" pageId="16" pageNumber="17">8.5</td>
|
||
<td box="[512,697,1134,1156]" gridcol="2" gridrow="20" pageId="16" pageNumber="17">20</td>
|
||
<td box="[733,865,1134,1156]" gridcol="3" gridrow="20" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1134,1155]" pageId="16" pageNumber="17">20</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1134,1156]" gridcol="4" gridrow="20" pageId="16" pageNumber="17">7</td>
|
||
<td box="[1101,1290,1134,1156]" gridcol="5" gridrow="20" pageId="16" pageNumber="17">20</td>
|
||
<td box="[1313,1508,1134,1156]" gridcol="6" gridrow="20" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,1134,1155]" pageId="16" pageNumber="17">19</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1171,1193]" gridrow="21" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1171,1193]" gridcol="0" gridrow="21" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,266,1171,1193]" collectionCode="NHMUK" pageId="16" pageNumber="17" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1171,1193]" gridcol="1" gridrow="21" pageId="16" pageNumber="17">7.4</td>
|
||
<td box="[512,697,1171,1193]" gridcol="2" gridrow="21" pageId="16" pageNumber="17">21</td>
|
||
<td box="[733,865,1171,1193]" gridcol="3" gridrow="21" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1171,1192]" pageId="16" pageNumber="17">21</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1171,1193]" gridcol="4" gridrow="21" pageId="16" pageNumber="17">3</td>
|
||
<td box="[1101,1290,1171,1193]" gridcol="5" gridrow="21" pageId="16" pageNumber="17">24</td>
|
||
<td box="[1313,1508,1171,1193]" gridcol="6" gridrow="21" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1171,1192]" pageId="16" pageNumber="17">23.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1209,1230]" gridrow="22" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1209,1230]" gridcol="0" gridrow="22" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,214,1209,1230]" collectionCode="MOR" pageId="16" pageNumber="17" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1209,1230]" gridcol="1" gridrow="22" pageId="16" pageNumber="17">7.1</td>
|
||
<td box="[512,697,1209,1230]" gridcol="2" gridrow="22" pageId="16" pageNumber="17">22</td>
|
||
<td box="[733,865,1209,1230]" gridcol="3" gridrow="22" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1209,1230]" pageId="16" pageNumber="17">22</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1209,1230]" gridcol="4" gridrow="22" pageId="16" pageNumber="17">2</td>
|
||
<td box="[1101,1290,1209,1230]" gridcol="5" gridrow="22" pageId="16" pageNumber="17">27</td>
|
||
<td box="[1313,1508,1209,1230]" gridcol="6" gridrow="22" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1209,1230]" pageId="16" pageNumber="17">26.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1246,1269]" gridrow="23" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1246,1269]" gridcol="0" gridrow="23" pageId="16" pageNumber="17">BMRP 2006.6.4</th>
|
||
<td box="[329,461,1246,1269]" gridcol="1" gridrow="23" pageId="16" pageNumber="17">6.8</td>
|
||
<td box="[512,697,1246,1269]" gridcol="2" gridrow="23" pageId="16" pageNumber="17">23</td>
|
||
<td box="[733,865,1246,1269]" gridcol="3" gridrow="23" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1246,1267]" pageId="16" pageNumber="17">23</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1246,1269]" gridcol="4" gridrow="23" pageId="16" pageNumber="17">89</td>
|
||
<td box="[1101,1290,1246,1269]" gridcol="5" gridrow="23" pageId="16" pageNumber="17">1</td>
|
||
<td box="[1313,1508,1246,1269]" gridcol="6" gridrow="23" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,1246,1267]" pageId="16" pageNumber="17">1.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1284,1306]" gridrow="24" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1284,1306]" gridcol="0" gridrow="24" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,299,1284,1306]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47763" pageId="16" pageNumber="17" specimenCode="AMNH FARB 5029">AMNH FARB 5029</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1284,1306]" gridcol="1" gridrow="24" pageId="16" pageNumber="17">6.6</td>
|
||
<td box="[512,697,1284,1306]" gridcol="2" gridrow="24" pageId="16" pageNumber="17">24</td>
|
||
<td box="[733,865,1284,1306]" gridcol="3" gridrow="24" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1284,1305]" pageId="16" pageNumber="17">24</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1284,1306]" gridcol="4" gridrow="24" pageId="16" pageNumber="17">10</td>
|
||
<td box="[1101,1290,1284,1306]" gridcol="5" gridrow="24" pageId="16" pageNumber="17">14</td>
|
||
<td box="[1313,1508,1284,1306]" gridcol="6" gridrow="24" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1284,1305]" pageId="16" pageNumber="17">13.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1322,1344]" gridrow="25" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1322,1344]" gridcol="0" gridrow="25" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,278,1322,1344]" collectionCode="NMMNH" pageId="16" pageNumber="17" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1322,1344]" gridcol="1" gridrow="25" pageId="16" pageNumber="17">5.8</td>
|
||
<td box="[512,697,1322,1344]" gridcol="2" gridrow="25" pageId="16" pageNumber="17">25</td>
|
||
<td box="[733,865,1322,1344]" gridcol="3" gridrow="25" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1322,1343]" pageId="16" pageNumber="17">25</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1322,1344]" gridcol="4" gridrow="25" pageId="16" pageNumber="17">1</td>
|
||
<td box="[1101,1290,1322,1344]" gridcol="5" gridrow="25" pageId="16" pageNumber="17">29</td>
|
||
<td box="[1313,1508,1322,1344]" gridcol="6" gridrow="25" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,1322,1343]" pageId="16" pageNumber="17">29</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1359,1381]" gridrow="26" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1359,1381]" gridcol="0" gridrow="26" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,224,1360,1381]" collectionCode="RSM" pageId="16" pageNumber="17" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1359,1381]" gridcol="1" gridrow="26" pageId="16" pageNumber="17">5.4</td>
|
||
<td box="[512,697,1359,1381]" gridcol="2" gridrow="26" pageId="16" pageNumber="17">26</td>
|
||
<td box="[733,865,1359,1381]" gridcol="3" gridrow="26" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1359,1380]" pageId="16" pageNumber="17">26</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1359,1381]" gridcol="4" gridrow="26" pageId="16" pageNumber="17">89</td>
|
||
<td box="[1101,1290,1359,1381]" gridcol="5" gridrow="26" pageId="16" pageNumber="17">2</td>
|
||
<td box="[1313,1508,1359,1381]" gridcol="6" gridrow="26" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,1359,1380]" pageId="16" pageNumber="17">1.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1397,1418]" gridrow="27" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1397,1418]" gridcol="0" gridrow="27" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,222,1397,1418]" collectionCode="DDM" pageId="16" pageNumber="17" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1397,1418]" gridcol="1" gridrow="27" pageId="16" pageNumber="17">3.7</td>
|
||
<td box="[512,697,1397,1418]" gridcol="2" gridrow="27" pageId="16" pageNumber="17">27</td>
|
||
<td box="[733,865,1397,1418]" gridcol="3" gridrow="27" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1397,1418]" pageId="16" pageNumber="17">27</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1397,1418]" gridcol="4" gridrow="27" pageId="16" pageNumber="17">0</td>
|
||
<td box="[1101,1290,1397,1418]" gridcol="5" gridrow="27" pageId="16" pageNumber="17">31</td>
|
||
<td box="[1313,1508,1397,1418]" gridcol="6" gridrow="27" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1397,1418]" pageId="16" pageNumber="17">30.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1434,1456]" gridrow="28" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1434,1456]" gridcol="0" gridrow="28" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,251,1435,1456]" collectionCode="UCMP" pageId="16" pageNumber="17" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1434,1456]" gridcol="1" gridrow="28" pageId="16" pageNumber="17">2.4</td>
|
||
<td box="[512,697,1434,1456]" gridcol="2" gridrow="28" pageId="16" pageNumber="17">28</td>
|
||
<td box="[733,865,1434,1456]" gridcol="3" gridrow="28" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1434,1455]" pageId="16" pageNumber="17">28</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1434,1456]" gridcol="4" gridrow="28" pageId="16" pageNumber="17">15</td>
|
||
<td box="[1101,1290,1434,1456]" gridcol="5" gridrow="28" pageId="16" pageNumber="17">10</td>
|
||
<td box="[1313,1508,1434,1456]" gridcol="6" gridrow="28" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,1434,1455]" pageId="16" pageNumber="17">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1472,1494]" gridrow="29" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1472,1494]" gridcol="0" gridrow="29" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,246,1472,1494]" collectionCode="UMNH" pageId="16" pageNumber="17" specimenCode="UMNH 11000">UMNH 11000</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1472,1494]" gridcol="1" gridrow="29" pageId="16" pageNumber="17">2.2</td>
|
||
<td box="[512,697,1472,1494]" gridcol="2" gridrow="29" pageId="16" pageNumber="17">29</td>
|
||
<td box="[733,865,1472,1494]" gridcol="3" gridrow="29" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,757,1472,1493]" pageId="16" pageNumber="17">29</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1472,1494]" gridcol="4" gridrow="29" pageId="16" pageNumber="17">15</td>
|
||
<td box="[1101,1290,1472,1494]" gridcol="5" gridrow="29" pageId="16" pageNumber="17">11</td>
|
||
<td box="[1313,1508,1472,1494]" gridcol="6" gridrow="29" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1343,1472,1493]" pageId="16" pageNumber="17">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1510,1532]" gridrow="30" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1510,1532]" gridcol="0" gridrow="30" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,299,1510,1532]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47787" pageId="16" pageNumber="17" specimenCode="AMNH FARB 5050">AMNH FARB 5050</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1510,1532]" gridcol="1" gridrow="30" pageId="16" pageNumber="17">1.8</td>
|
||
<td box="[512,697,1510,1532]" gridcol="2" gridrow="30" pageId="16" pageNumber="17">30</td>
|
||
<td box="[733,865,1510,1532]" gridcol="3" gridrow="30" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,775,1510,1531]" pageId="16" pageNumber="17">30.5</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1510,1532]" gridcol="4" gridrow="30" pageId="16" pageNumber="17">2</td>
|
||
<td box="[1101,1290,1510,1532]" gridcol="5" gridrow="30" pageId="16" pageNumber="17">28</td>
|
||
<td box="[1313,1508,1510,1532]" gridcol="6" gridrow="30" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1355,1510,1531]" pageId="16" pageNumber="17">26.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1508,1547,1569]" gridrow="31" pageId="16" pageNumber="17">
|
||
<th box="[108,301,1547,1569]" gridcol="0" gridrow="31" pageId="16" pageNumber="17">
|
||
<materialsCitation box="[108,301,1547,1569]" collectionCode="TMP" pageId="16" pageNumber="17" specimenCode="TMP 1981.006.0001">TMP 1981.012.0001</materialsCitation>
|
||
</th>
|
||
<td box="[329,461,1547,1569]" gridcol="1" gridrow="31" pageId="16" pageNumber="17">1.8</td>
|
||
<td box="[512,697,1547,1569]" gridcol="2" gridrow="31" pageId="16" pageNumber="17">31</td>
|
||
<td box="[733,865,1547,1569]" gridcol="3" gridrow="31" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[733,775,1547,1568]" pageId="16" pageNumber="17">30.5</emphasis>
|
||
</td>
|
||
<td box="[887,1083,1547,1569]" gridcol="4" gridrow="31" pageId="16" pageNumber="17">9</td>
|
||
<td box="[1101,1290,1547,1569]" gridcol="5" gridrow="31" pageId="16" pageNumber="17">17</td>
|
||
<td box="[1313,1508,1547,1569]" gridcol="6" gridrow="31" pageId="16" pageNumber="17">
|
||
<emphasis bold="true" box="[1313,1337,1547,1568]" pageId="16" pageNumber="17">16</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824978" ID-Zenodo-Dep="5824978" httpUri="https://zenodo.org/record/5824978/files/figure.png" pageId="17" pageNumber="18" startId="17.[524,589,657,679]" targetBox="[473,1323,220,638]" targetPageId="17">
|
||
<paragraph blockId="17.[524,1515,657,855]" pageId="17" pageNumber="18">
|
||
<emphasis bold="true" pageId="17" pageNumber="18">
|
||
Figure 3 Scatterplot showing the noncongruence in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1043,1228,658,679]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1043,1228,658,679]" italics="true" pageId="17" pageNumber="18">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
between the completeness of specimens (i.e., number of characters scored) and the number of synontomorphies at each corresponding node.
|
||
</emphasis>
|
||
Per cent completeness (decreasing away from the origin) and the number of synontomorphies supporting the corresponding node (decreasing away from the origin) have been converted to ranks. A Spearman correlation test on these data results in a nonsignificant correlation coefficient; ergo, the number of synontomorphies at an internode is not an artifact of specimen completeness. Full-size DOI: 10.7717/peerj.9192/fig-3
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824980" ID-Zenodo-Dep="5824980" httpUri="https://zenodo.org/record/5824980/files/figure.png" pageId="17" pageNumber="18" startId="17.[524,589,1366,1388]" targetBox="[500,1530,910,1333]" targetPageId="17">
|
||
<paragraph blockId="17.[524,1515,1366,1535]" pageId="17" pageNumber="18">
|
||
<emphasis bold="true" pageId="17" pageNumber="18">
|
||
Figure 4 Frequency distribution of unambiguously optimized synontomorphies during the growth of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[550,736,1396,1417]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[550,736,1396,1417]" italics="true" pageId="17" pageNumber="18">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages (corresponding to the numbered nodes of the ontogram in Fig. 2) are along the
|
||
<emphasis box="[653,664,1426,1447]" italics="true" pageId="17" pageNumber="18">x</emphasis>
|
||
-axis and the number of changes are along the
|
||
<emphasis box="[1109,1119,1426,1447]" italics="true" pageId="17" pageNumber="18">y</emphasis>
|
||
-axis. The greatest number of changes are seen in the transition from large juvenile to subadult, or, from growth stage 5
|
||
<emphasis box="[1243,1255,1454,1476]" italics="true" pageId="17" pageNumber="18">–</emphasis>
|
||
6; the high concentration of change between these growth categories is evidence that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1066,1121,1484,1505]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1066,1121,1484,1505]" italics="true" pageId="17" pageNumber="18">T. rex</emphasis>
|
||
</taxonomicName>
|
||
ontogeny is metamorphic (sensu
|
||
<bibRefCitation author="Rose CS & Reiss JO" editor="Hanken H & Hall BK" journalOrPublisher="Chicago: University of Chicago Press" pageId="17" pageNumber="18" refId="ref55796" refString="Rose CS, Reiss JO. 1993. Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms. In: Hanken H, Hall BK, eds. The Skull: Development. Vol. 1. Chicago: University of Chicago Press." title="Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms" type="book" volumeTitle="The Skull: Development. Vol. 1" year="1993">
|
||
<emphasis italics="true" pageId="17" pageNumber="18">Rose & Reiss, 1993</emphasis>
|
||
</bibRefCitation>
|
||
). Full-size DOI: 10.7717/peerj.9192/fig-4
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="17.[498,1542,1582,1928]" lastBlockId="19.[498,1542,975,1918]" lastPageId="19" lastPageNumber="20" pageId="17" pageNumber="18">
|
||
A comparison of the frequency of cranial and postcranial changes shows that cranial synontomorphies are the most frequent (
|
||
<figureCitation box="[984,1049,1622,1648]" captionStart="Figure 6" captionStartId="18.[524,589,1471,1493]" captionTargetBox="[498,1541,1003,1458]" captionTargetId="figure-71@18.[497,1165,1002,1445]" captionTargetPageId="18" captionText="Figure 6 Comparison of the frequency distributions of cranial and postcranial changes in the growth series of Tyrannosaurus rex. The growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Cranial changes are shown in solid bars; postcranial chanages are shown in hollow bars. Cranial and postcranial changes tend to follow the same overall pattern although postcranial changes are exceeded by cranial changes, except at growth stages 7, 15, and 16. The relatively late occurrence of postcranial changes (at growth stage 6) is an artifact of the absence of postcranial material among the least mature specimens in the sample. Full-size DOI: 10.7717/peerj.9192/fig-6" figureDoi="http://doi.org/10.5281/zenodo.5824984" httpUri="https://zenodo.org/record/5824984/files/figure.png" pageId="17" pageNumber="18">Fig. 6</figureCitation>
|
||
). Both sets of changes follow the same general pattern; cranial changes are dominant in terms of number and the pattern of the frequency distribution, whereas postcranial changes do not always occur. Cranial and postcranial changes are most frequent at growth stage 6, and thereafter they show less than 22 changes per growth stage. This indicates that once the adult morphotype (i.e., tall skull, inflated antorbital sinuses) is achieved, the rate of change, as shown by the number of changes per growth stage, greatly decreases. During adulthood, cranial changes generally outnumber postcranial changes, which tend to cease altogether. Cranial changes are nearly continuous and have five peaks, whereas only three are seen postcranially that are often preceded and followed by one or more growth stages of no change. The absence of postcranial changes in growth stage 5 is an artifact of the least mature specimens lacking postcranial material. The abundance of postcranial changes at growth stage 7 is an artifact of the exemplar specimen having an incomplete skull and a relatively complete hindlimb. In contrast, the absence of postcranial change seen from growth stage 9
|
||
<emphasis box="[1466,1481,1015,1040]" italics="true" pageId="19" pageNumber="20">–</emphasis>
|
||
12 reflects a signal of quiescence given that several specimens in that interval do include postcranial bones.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824982" ID-Zenodo-Dep="5824982" httpUri="https://zenodo.org/record/5824982/files/figure.png" pageId="18" pageNumber="19" startId="18.[524,589,692,714]" targetBox="[498,1541,234,679]" targetPageId="18">
|
||
<paragraph blockId="18.[524,1516,692,948]" pageId="18" pageNumber="19">
|
||
<emphasis bold="true" pageId="18" pageNumber="19">
|
||
Figure 5 Comparison of the frequency distributions of phylogenetic and nonphylogenetic synontomorphies in the ontogeny of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[911,1100,722,743]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[911,1100,722,743]" italics="true" pageId="18" pageNumber="19">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stage is along the
|
||
<emphasis box="[1383,1394,722,743]" italics="true" pageId="18" pageNumber="19">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and number of synontomorphies is along the
|
||
<emphasis box="[524,534,781,802]" italics="true" pageId="18" pageNumber="19">y</emphasis>
|
||
-axis. Phylogenetic characters are in solid bars; nonphylogenetic characters are in hollow bars. The frequency distributions of both sets of data follow the same general pattern, aside from the flatter distribution of the phylogenetic synontomorphies relative to the nonphylogenetic synontomorphies and the reversed pattern seen at growth stages 7 and 8. Both types of changes occur throughout the lifespan of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,579,898,919]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[524,579,898,919]" italics="true" pageId="18" pageNumber="19">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, indicating that ontogeny is not strictly congruent with phylogeny.
|
||
</paragraph>
|
||
<paragraph blockId="18.[524,1516,692,948]" box="[1110,1515,926,948]" pageId="18" pageNumber="19">Full-size DOI: 10.7717/peerj.9192/fig-5</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824984" ID-Zenodo-Dep="5824984" httpUri="https://zenodo.org/record/5824984/files/figure.png" pageId="18" pageNumber="19" startId="18.[524,589,1471,1493]" targetBox="[498,1541,1003,1458]" targetPageId="18">
|
||
<paragraph blockId="18.[524,1515,1471,1698]" pageId="18" pageNumber="19">
|
||
<emphasis bold="true" pageId="18" pageNumber="19">
|
||
Figure 6 Comparison of the frequency distributions of cranial and postcranial changes in the growth series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[614,802,1501,1522]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[614,802,1501,1522]" italics="true" pageId="18" pageNumber="19">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The growth stages are along the
|
||
<emphasis box="[1134,1145,1501,1522]" italics="true" pageId="18" pageNumber="19">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[915,925,1530,1551]" italics="true" pageId="18" pageNumber="19">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. Cranial changes are shown in solid bars; postcranial chanages are shown in hollow bars. Cranial and postcranial changes tend to follow the same overall pattern although postcranial changes are exceeded by cranial changes, except at growth stages 7, 15, and 16. The relatively late occurrence of postcranial changes (at growth stage 6) is an artifact of the absence of postcranial material among the least mature specimens in the sample. Full-size DOI: 10.7717/peerj.9192/fig-6
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824986" ID-Zenodo-Dep="5824986" httpUri="https://zenodo.org/record/5824986/files/figure.png" pageId="19" pageNumber="20" startId="19.[524,589,693,715]" targetBox="[501,1534,234,662]" targetPageId="19">
|
||
<paragraph blockId="19.[524,1515,693,920]" pageId="19" pageNumber="20">
|
||
<emphasis bold="true" pageId="19" pageNumber="20">
|
||
Figure 7 Comparison of the frequency distribution of synontomorphies of the cranium with that of the mandibular ramus in the ontogeny of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[966,1155,723,744]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[966,1155,723,744]" italics="true" pageId="19" pageNumber="20">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages are along the
|
||
<emphasis box="[1459,1470,723,744]" italics="true" pageId="19" pageNumber="20">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[1268,1278,752,773]" italics="true" pageId="19" pageNumber="20">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. Skull changes are shown with solid bars; mandible changes are shown with hollow bars. Although a greater number of changes is seen in the cranium than in the mandibular ramus, the lower jaw completes its early phase of changes (stage 5) before the cranium (stage 6). Thereafter, the pattern of mandibular changes is generally congruent with the cranium.
|
||
</paragraph>
|
||
<paragraph blockId="19.[524,1515,693,920]" box="[1110,1515,898,920]" pageId="19" pageNumber="20">Full-size DOI: 10.7717/peerj.9192/fig-7</paragraph>
|
||
</caption>
|
||
<paragraph blockId="19.[498,1542,975,1918]" pageId="19" pageNumber="20">
|
||
When the frequencies of changes to the cranium and mandible are compared, both follow the same overall pattern (
|
||
<figureCitation box="[872,935,1174,1200]" captionStart="Figure 7" captionStartId="19.[524,589,693,715]" captionTargetBox="[501,1534,234,662]" captionTargetId="figure-324@19.[497,1131,233,666]" captionTargetPageId="19" captionText="Figure 7 Comparison of the frequency distribution of synontomorphies of the cranium with that of the mandibular ramus in the ontogeny of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Skull changes are shown with solid bars; mandible changes are shown with hollow bars. Although a greater number of changes is seen in the cranium than in the mandibular ramus, the lower jaw completes its early phase of changes (stage 5) before the cranium (stage 6). Thereafter, the pattern of mandibular changes is generally congruent with the cranium. Full-size DOI: 10.7717/peerj.9192/fig-7" figureDoi="http://doi.org/10.5281/zenodo.5824986" httpUri="https://zenodo.org/record/5824986/files/figure.png" pageId="19" pageNumber="20">Fig. 7</figureCitation>
|
||
), but the cranial changes generally outnumber those of the mandible. Mandibular changes occur in a series of three low peaks (i.e., less than 8 synontomorphies per growth stage) throughout ontogeny; the highest number of changes (7) occurs early in the 5th growth stage, but this trend does not continue into the sustained and extremely high number of changes seen in the cranium at the 6th stage. This difference results from the absence of mandibular bones in the sample of subadults at the fifth stage. In contrast, the exemplar of the seventh growth stage,
|
||
<materialsCitation box="[1318,1479,1413,1440]" collectionCode="LACM" pageId="19" pageNumber="20" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
, is represented by an incomplete mandibular ramus, but, notably, it does not result in mandibular synontomorphies at that growth stage; presumably those changes occurred earlier, at growth stage 5. The absence of changes at the 7th growth stage indicates that the completion of the progression of changes in the mandibular ramus, which produces the dorsoventrally deep skull frame, precedes that of the cranium. In adulthood, the number of changes in the cranium exceeds that of the mandible or, on occasion, mandibular changes exceed cranial changes at growth stages where the cephalic skeleton is represented solely by mandibular bones (
|
||
<materialsCitation box="[982,1180,1732,1758]" collectionCode="NHMUK" pageId="19" pageNumber="20" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
) or very few cranial bones (
|
||
<materialsCitation box="[506,722,1772,1799]" collectionCode="NMMNH" pageId="19" pageNumber="20" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="19.[498,1542,975,1918]" lastBlockId="20.[498,1542,947,1930]" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
|
||
Changes to the skull and jaws were examined by anatomical domain, which includes discrete regions of the skull such as bony structures associated with the antorbital air sac system or aggregates of functional structures, such as joint surfaces. For ease of comparison, the domains were separated into two groups: pneumatic systems on the one hand, and apneumatic features on the other. The pneumatic systems include the antorbital sinus system (invades the snout), tympanic sinus system (invades the lateral surface of the braincase), median pharyngeal sinus system (invades the sphenoid rostrum and basicranium), and the cervical air sac system (invades the occiput). In descending order, the most frequent pneumatic changes are associated with the antorbital, cervical (=subcondylar) and tympanic sinus systems, and the median pharyngeal sinus system (
|
||
<figureCitation box="[508,572,1226,1252]" captionStart="Figure 8" captionStartId="20.[524,589,694,716]" captionTargetBox="[498,1541,235,681]" captionTargetId="figure-333@20.[497,1131,233,668]" captionTargetPageId="20" captionText="Figure 8 The frequency distribution of synontomorphies by cephalic pneumatic system in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Changes to the antorbital sinus system are dominant over others and are sustained though growth, in contrast to the other systems that occur in adulthood and are transient in occurrence. aosin, antorbital sinus system; pharyn, pharyngeal sinus system; subcon, subcondylar sinus system; tympcav, tympanic cavity. Full-size DOI: 10.7717/peerj.9192/fig-8" figureDoi="http://doi.org/10.5281/zenodo.5824988" httpUri="https://zenodo.org/record/5824988/files/figure.png" pageId="20" pageNumber="21">Fig. 8</figureCitation>
|
||
). Changes associated with the antorbital sinus occur throughout ontogeny, whereas cervical changes begin in adulthood and continue almost to senescence; in contrast, median pharyngeal changes are limited to growth stage 12, at the end of the young adult growth category. Finally, tympanic changes are limited to growth stages 9
|
||
<emphasis box="[1367,1382,1346,1371]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
12, during young adulthood.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824988" ID-Zenodo-Dep="5824988" httpUri="https://zenodo.org/record/5824988/files/figure.png" pageId="20" pageNumber="21" startId="20.[524,589,694,716]" targetBox="[498,1541,235,681]" targetPageId="20">
|
||
<paragraph blockId="20.[524,1516,694,892]" pageId="20" pageNumber="21">
|
||
<emphasis bold="true" pageId="20" pageNumber="21">
|
||
Figure 8 The frequency distribution of synontomorphies by cephalic pneumatic system in the growth series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[694,882,724,745]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[694,882,724,745]" italics="true" pageId="20" pageNumber="21">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages are along the
|
||
<emphasis box="[1178,1189,724,745]" italics="true" pageId="20" pageNumber="21">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[985,995,754,775]" italics="true" pageId="20" pageNumber="21">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. Changes to the antorbital sinus system are dominant over others and are sustained though growth, in contrast to the other systems that occur in adulthood and are transient in occurrence. aosin, antorbital sinus system; pharyn, pharyngeal sinus system; subcon, subcondylar sinus system; tympcav, tympanic cavity. Full-size DOI: 10.7717/peerj.9192/fig-8
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="20.[498,1542,947,1930]" pageId="20" pageNumber="21">
|
||
Apneumatic domains include joint surfaces, muscle scars, the subcutaneous surface, neurovasculature, the occiput, dentition, and the skull frame. The skull frame sees the most changes throughout the growth series (
|
||
<figureCitation box="[960,1025,1505,1531]" captionStart="Figure 9" captionStartId="21.[524,589,775,797]" captionTargetBox="[503,1533,234,742]" captionTargetId="figure-102@21.[497,1237,233,749]" captionTargetPageId="21" captionText="Figure 9 The frequency distribution of synontomorphies by apneumatic anatomical domain in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Changes to the skull frame are dominant over others and all are sustained throughout growth, aside from the dentition and cervical occiput. crv occ, cervical occiput; dntn, dentition; dtfo, dorsotemporal fossa; jnt srfc, joint surfaces; mscl scrs, muscle scars; nrvsc, neurovasculature; sbct srfc, subcutaneous surface; skl frm, skull frame. Full-size DOI: 10.7717/peerj.9192/fig-9" figureDoi="http://doi.org/10.5281/zenodo.5824990" httpUri="https://zenodo.org/record/5824990/files/figure.png" pageId="20" pageNumber="21">Fig. 9</figureCitation>
|
||
). Change to all other domains are low in frequency (i.e., fewer than 10 changes per growth stage), but most of them also change throughout growth, including joint surfaces (growth stage 4
|
||
<emphasis box="[1195,1210,1585,1610]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
21), the dorsotemporal fossa (growth stage 4
|
||
<emphasis box="[680,695,1625,1650]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
21), neurovasculature (growth stage 2
|
||
<emphasis box="[1139,1154,1625,1650]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
21), muscle scars (growth stage 5
|
||
<emphasis box="[512,527,1665,1690]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
21), and the subcutaneous surface (growth stage 5
|
||
<emphasis box="[1111,1126,1665,1690]" italics="true" pageId="20" pageNumber="21">–</emphasis>
|
||
21). A limited pattern is seen in the dentition where changes occur from the 5th to 17th growth stages. In contrast, nonmuscular and apneumatic changes to the occiput are limited to the young adult growth category (growth stage 9).
|
||
</paragraph>
|
||
<paragraph blockId="20.[498,1542,947,1930]" lastBlockId="21.[498,1542,1683,1909]" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21">
|
||
In the postcranial skeleton, the greatest number of changes (more than five) are seen in the transition from the subadult to the young adult growth categories (
|
||
<figureCitation box="[1306,1382,1863,1890]" captionStart="Figure 10" captionStartId="21.[524,589,1489,1511]" captionTargetBox="[503,1531,1030,1457]" captionTargetId="figure-217@21.[497,1128,1026,1462]" captionTargetPageId="21" captionText="Figure 10 The frequency distribution of postcranial synontomorphies in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Changes to the appendicular skeleton dominate in the transition between juvenile and subadult, whereas changes to the pelvic girdle and axial skeleton occur late in adulthood. Full-size DOI: 10.7717/peerj.9192/fig-10" figureDoi="http://doi.org/10.5281/zenodo.5824992" httpUri="https://zenodo.org/record/5824992/files/figure.png" pageId="20" pageNumber="21">Fig. 10</figureCitation>
|
||
) that include, in descending order, the pes, fibula, scapula, coracoid, and humerus. Thereafter, postcranial changes effectively cease until three clusters of changes in adulthood, at the 14th and 15th stages, the 18th and 19th stages, and a final set at the 21st growth stage.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824990" ID-Zenodo-Dep="5824990" httpUri="https://zenodo.org/record/5824990/files/figure.png" pageId="21" pageNumber="22" startId="21.[524,589,775,797]" targetBox="[503,1533,234,742]" targetPageId="21">
|
||
<paragraph blockId="21.[524,1516,775,973]" pageId="21" pageNumber="22">
|
||
<emphasis bold="true" pageId="21" pageNumber="22">
|
||
Figure 9 The frequency distribution of synontomorphies by apneumatic anatomical domain in the growth series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[684,869,805,826]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[684,869,805,826]" italics="true" pageId="21" pageNumber="22">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages are along the
|
||
<emphasis box="[1146,1157,805,826]" italics="true" pageId="21" pageNumber="22">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[909,919,834,855]" italics="true" pageId="21" pageNumber="22">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. Changes to the skull frame are dominant over others and all are sustained throughout growth, aside from the dentition and cervical occiput. crv occ, cervical occiput; dntn, dentition; dtfo, dorsotemporal fossa; jnt srfc, joint surfaces; mscl scrs, muscle scars; nrvsc, neurovasculature; sbct srfc, subcutaneous surface; skl frm, skull frame. Full-size DOI: 10.7717/peerj.9192/fig-9
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824992" ID-Zenodo-Dep="5824992" httpUri="https://zenodo.org/record/5824992/files/figure.png" pageId="21" pageNumber="22" startId="21.[524,589,1489,1511]" targetBox="[503,1531,1030,1457]" targetPageId="21">
|
||
<paragraph blockId="21.[524,1515,1489,1628]" pageId="21" pageNumber="22">
|
||
<emphasis bold="true" pageId="21" pageNumber="22">
|
||
Figure 10 The frequency distribution of postcranial synontomorphies in the growth series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,711,1518,1539]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[524,711,1518,1539]" italics="true" pageId="21" pageNumber="22">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages are along the
|
||
<emphasis box="[1005,1016,1519,1540]" italics="true" pageId="21" pageNumber="22">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[830,840,1548,1569]" italics="true" pageId="21" pageNumber="22">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. Changes to the appendicular skeleton dominate in the transition between juvenile and subadult, whereas changes to the pelvic girdle and axial skeleton occur late in adulthood. Full-size DOI: 10.7717/peerj.9192/fig-10
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="21.[498,1542,1683,1909]" lastBlockId="22.[498,1521,1443,1549]" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22">Early changes (juvenile to young adult categories) happen to the pectoral girdle and pes (6th and 7th growth stages), the humerus (6th growth stage), and the fibula (8th growth stage). Later changes (adult to senescent adult categories) are seen in the axial column (15th, 19th, and 21st growth stages), pelvic girdle (14th and 18th growth stages), and the fibula (18th and 19th growth stages). This pattern suggests a relatively instantaneous transition in the pectoral girdle and limb, and the pes; in contrast, changes to the axial column, and to the pelvic girdle and limb, are relatively sustained throughout growth.</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824994" ID-Zenodo-Dep="5824994" httpUri="https://zenodo.org/record/5824994/files/figure.png" pageId="22" pageNumber="23" startId="22.[524,589,695,717]" targetBox="[502,1535,236,665]" targetPageId="22">
|
||
<paragraph blockId="22.[524,1515,695,893]" pageId="22" pageNumber="23">
|
||
<emphasis bold="true" pageId="22" pageNumber="23">
|
||
Figure 11 The frequency distribution of changes to the craniomandibular functional modules (sensu
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[524,747,724,746]" journalOrPublisher="Scientific Reports" pageId="22" pageNumber="23" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis bold="true" box="[524,747,724,746]" italics="true" pageId="22" pageNumber="23">Werneburg et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
) in the growth series of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[989,1175,725,746]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[989,1175,725,746]" italics="true" pageId="22" pageNumber="23">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages are along the
|
||
<emphasis box="[1459,1470,726,747]" italics="true" pageId="22" pageNumber="23">x</emphasis>
|
||
-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the
|
||
<emphasis box="[1268,1278,755,776]" italics="true" pageId="22" pageNumber="23">y</emphasis>
|
||
-axis corresponds to the number of synontomorphies. The onset of the changes to the skull roof, snout, mandibular ramus, and suspensorium modules occur early in growth, whereas the onset of changes to the parietal and braincase occur in adulthood. Changes continue throughout growth in all domains, aside from those to the parietal that cease at growth stage 14. Full-size DOI: 10.7717/peerj.9192/fig-11
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="22" pageNumber="23" startId="22.[117,172,963,984]" targetBox="[108,1525,1064,1353]" targetIsTable="true" targetPageId="22">
|
||
<paragraph blockId="22.[117,1515,963,1044]" pageId="22" pageNumber="23">
|
||
<emphasis bold="true" pageId="22" pageNumber="23">
|
||
Table 4 Summary of synontomorphies and individual variation in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[786,972,963,984]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[786,972,963,984]" italics="true" pageId="22" pageNumber="23">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
organized by growth category and functional modules of the skull and jaws.
|
||
</emphasis>
|
||
Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis italics="true" pageId="22" pageNumber="23">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
, organized by functional modules (sensu
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[550,767,1021,1043]" journalOrPublisher="Scientific Reports" pageId="22" pageNumber="23" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis box="[550,767,1021,1043]" italics="true" pageId="22" pageNumber="23">Werneburg et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
). Individual variation is shown in parentheses.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="22" pageNumber="23">
|
||
<table box="[108,1525,1064,1353]" gridcols="8" gridrows="8" pageId="22" pageNumber="23">
|
||
<tr box="[108,1525,1064,1086]" gridrow="0" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1064,1086]" gridcol="0" gridrow="0" pageId="22" pageNumber="23">Growth category</th>
|
||
<th box="[346,446,1064,1086]" gridcol="1" gridrow="0" pageId="22" pageNumber="23">Skull roof</th>
|
||
<th box="[517,664,1064,1086]" gridcol="2" gridrow="0" pageId="22" pageNumber="23">Snout + palate</th>
|
||
<th box="[736,813,1064,1086]" gridcol="3" gridrow="0" pageId="22" pageNumber="23">Parietal</th>
|
||
<th box="[884,1026,1064,1086]" gridcol="4" gridrow="0" pageId="22" pageNumber="23">Suspensorium</th>
|
||
<th box="[1098,1194,1064,1086]" gridcol="5" gridrow="0" pageId="22" pageNumber="23">Braincase</th>
|
||
<th box="[1266,1361,1064,1086]" gridcol="6" gridrow="0" pageId="22" pageNumber="23">Mandible</th>
|
||
<th box="[1432,1525,1064,1086]" gridcol="7" gridrow="0" pageId="22" pageNumber="23">
|
||
<emphasis bold="true" box="[1432,1485,1064,1085]" pageId="22" pageNumber="23">Total</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1525,1105,1127]" gridrow="1" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1105,1127]" gridcol="0" gridrow="1" pageId="22" pageNumber="23">Small juvenile</th>
|
||
<td box="[346,446,1105,1127]" gridcol="1" gridrow="1" pageId="22" pageNumber="23">3 (3)</td>
|
||
<td box="[517,664,1105,1127]" gridcol="2" gridrow="1" pageId="22" pageNumber="23">0 (1)</td>
|
||
<td box="[736,813,1105,1127]" gridcol="3" gridrow="1" pageId="22" pageNumber="23">0 (2)</td>
|
||
<td box="[884,1026,1105,1127]" gridcol="4" gridrow="1" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[1098,1194,1105,1127]" gridcol="5" gridrow="1" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[1266,1361,1105,1127]" gridcol="6" gridrow="1" pageId="22" pageNumber="23">2 (0)</td>
|
||
<td box="[1432,1525,1105,1127]" gridcol="7" gridrow="1" pageId="22" pageNumber="23">5 (6)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1142,1165]" gridrow="2" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1142,1165]" gridcol="0" gridrow="2" pageId="22" pageNumber="23">Large juvenile</th>
|
||
<td box="[346,446,1142,1165]" gridcol="1" gridrow="2" pageId="22" pageNumber="23">15 (6)</td>
|
||
<td box="[517,664,1142,1165]" gridcol="2" gridrow="2" pageId="22" pageNumber="23">30 (5)</td>
|
||
<td box="[736,813,1142,1165]" gridcol="3" gridrow="2" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[884,1026,1142,1165]" gridcol="4" gridrow="2" pageId="22" pageNumber="23">1 (0)</td>
|
||
<td box="[1098,1194,1142,1165]" gridcol="5" gridrow="2" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[1266,1361,1142,1165]" gridcol="6" gridrow="2" pageId="22" pageNumber="23">7 (3)</td>
|
||
<td box="[1432,1525,1142,1165]" gridcol="7" gridrow="2" pageId="22" pageNumber="23">53 (14)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1180,1202]" gridrow="3" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1180,1202]" gridcol="0" gridrow="3" pageId="22" pageNumber="23">Subadult</th>
|
||
<td box="[346,446,1180,1202]" gridcol="1" gridrow="3" pageId="22" pageNumber="23">58 (2)</td>
|
||
<td box="[517,664,1180,1202]" gridcol="2" gridrow="3" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[736,813,1180,1202]" gridcol="3" gridrow="3" pageId="22" pageNumber="23">0 (2)</td>
|
||
<td box="[884,1026,1180,1202]" gridcol="4" gridrow="3" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[1098,1194,1180,1202]" gridcol="5" gridrow="3" pageId="22" pageNumber="23">0 (0)</td>
|
||
<td box="[1266,1361,1180,1202]" gridcol="6" gridrow="3" pageId="22" pageNumber="23">0 (1)</td>
|
||
<td box="[1432,1525,1180,1202]" gridcol="7" gridrow="3" pageId="22" pageNumber="23">58 (5)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1218,1240]" gridrow="4" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1218,1240]" gridcol="0" gridrow="4" pageId="22" pageNumber="23">Young adult</th>
|
||
<td box="[346,446,1218,1240]" gridcol="1" gridrow="4" pageId="22" pageNumber="23">17 (38)</td>
|
||
<td box="[517,664,1218,1240]" gridcol="2" gridrow="4" pageId="22" pageNumber="23">13 (23)</td>
|
||
<td box="[736,813,1218,1240]" gridcol="3" gridrow="4" pageId="22" pageNumber="23">11 (10)</td>
|
||
<td box="[884,1026,1218,1240]" gridcol="4" gridrow="4" pageId="22" pageNumber="23">5 (8)</td>
|
||
<td box="[1098,1194,1218,1240]" gridcol="5" gridrow="4" pageId="22" pageNumber="23">14 (8)</td>
|
||
<td box="[1266,1361,1218,1240]" gridcol="6" gridrow="4" pageId="22" pageNumber="23">7 (11)</td>
|
||
<td box="[1432,1525,1218,1240]" gridcol="7" gridrow="4" pageId="22" pageNumber="23">67 (98)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1255,1278]" gridrow="5" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1255,1278]" gridcol="0" gridrow="5" pageId="22" pageNumber="23">Adult</th>
|
||
<td box="[346,446,1255,1278]" gridcol="1" gridrow="5" pageId="22" pageNumber="23">14 (51)</td>
|
||
<td box="[517,664,1255,1278]" gridcol="2" gridrow="5" pageId="22" pageNumber="23">15 (28)</td>
|
||
<td box="[736,813,1255,1278]" gridcol="3" gridrow="5" pageId="22" pageNumber="23">1 (9)</td>
|
||
<td box="[884,1026,1255,1278]" gridcol="4" gridrow="5" pageId="22" pageNumber="23">7 (15)</td>
|
||
<td box="[1098,1194,1255,1278]" gridcol="5" gridrow="5" pageId="22" pageNumber="23">8 (21)</td>
|
||
<td box="[1266,1361,1255,1278]" gridcol="6" gridrow="5" pageId="22" pageNumber="23">11 (27)</td>
|
||
<td box="[1432,1525,1255,1278]" gridcol="7" gridrow="5" pageId="22" pageNumber="23">56 (151)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1293,1315]" gridrow="6" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1293,1315]" gridcol="0" gridrow="6" pageId="22" pageNumber="23">Senescent adult</th>
|
||
<td box="[346,446,1293,1315]" gridcol="1" gridrow="6" pageId="22" pageNumber="23">10 (n/a)</td>
|
||
<td box="[517,664,1293,1315]" gridcol="2" gridrow="6" pageId="22" pageNumber="23">2 (n/a)</td>
|
||
<td box="[736,813,1293,1315]" gridcol="3" gridrow="6" pageId="22" pageNumber="23">0 (n/a)</td>
|
||
<td box="[884,1026,1293,1315]" gridcol="4" gridrow="6" pageId="22" pageNumber="23">2 (n/a)</td>
|
||
<td box="[1098,1194,1293,1315]" gridcol="5" gridrow="6" pageId="22" pageNumber="23">1 (n/a)</td>
|
||
<td box="[1266,1361,1293,1315]" gridcol="6" gridrow="6" pageId="22" pageNumber="23">0 (n/a)</td>
|
||
<td box="[1432,1525,1293,1315]" gridcol="7" gridrow="6" pageId="22" pageNumber="23">15 (n/a)</td>
|
||
</tr>
|
||
<tr box="[108,1525,1330,1353]" gridrow="7" pageId="22" pageNumber="23">
|
||
<th box="[108,275,1330,1353]" gridcol="0" gridrow="7" pageId="22" pageNumber="23">Total</th>
|
||
<td box="[346,446,1330,1353]" gridcol="1" gridrow="7" pageId="22" pageNumber="23">117 (100)</td>
|
||
<td box="[517,664,1330,1353]" gridcol="2" gridrow="7" pageId="22" pageNumber="23">60 (57)</td>
|
||
<td box="[736,813,1330,1353]" gridcol="3" gridrow="7" pageId="22" pageNumber="23">12 (23)</td>
|
||
<td box="[884,1026,1330,1353]" gridcol="4" gridrow="7" pageId="22" pageNumber="23">13 (23)</td>
|
||
<td box="[1098,1194,1330,1353]" gridcol="5" gridrow="7" pageId="22" pageNumber="23">25 (29)</td>
|
||
<td box="[1266,1361,1330,1353]" gridcol="6" gridrow="7" pageId="22" pageNumber="23">27 (42)</td>
|
||
<td box="[1432,1525,1330,1353]" gridcol="7" gridrow="7" pageId="22" pageNumber="23">254 (274)</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="22.[498,1542,1585,1935]" box="[498,873,1585,1613]" pageId="22" pageNumber="23">
|
||
<heading bold="true" box="[498,873,1585,1613]" fontSize="11" level="3" pageId="22" pageNumber="23" reason="6">
|
||
<emphasis bold="true" box="[498,873,1585,1613]" italics="true" pageId="22" pageNumber="23">Craniomandibular modules</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="22.[498,1542,1585,1935]" lastBlockId="23.[498,1542,234,938]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
|
||
Recent work has found that the skull of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[968,1035,1630,1655]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[968,1035,1630,1655]" italics="true" pageId="22" pageNumber="23">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is organized into six modules of functional integration (
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[645,911,1669,1695]" journalOrPublisher="Scientific Reports" pageId="22" pageNumber="23" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis box="[645,911,1669,1695]" italics="true" pageId="22" pageNumber="23">Werneburg et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
). The growth series obtained here shows that the modules experience different amounts of ontogenetic change (
|
||
<figureCitation box="[1234,1313,1709,1735]" captionStart="Figure 11" captionStartId="22.[524,589,695,717]" captionTargetBox="[502,1535,236,665]" captionTargetId="figure-171@22.[497,1130,233,669]" captionTargetPageId="22" captionText="Figure 11 The frequency distribution of changes to the craniomandibular functional modules (sensu Werneburg et al., 2019) in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. The onset of the changes to the skull roof, snout, mandibular ramus, and suspensorium modules occur early in growth, whereas the onset of changes to the parietal and braincase occur in adulthood. Changes continue throughout growth in all domains, aside from those to the parietal that cease at growth stage 14. Full-size DOI: 10.7717/peerj.9192/fig-11" figureDoi="http://doi.org/10.5281/zenodo.5824994" httpUri="https://zenodo.org/record/5824994/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
|
||
;
|
||
<tableCitation box="[1328,1416,1709,1735]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="22" pageNumber="23" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
): in descending order, the dorsum of the snout, circumorbital bones, and frontals; the sides of the snout and palate; the mandibular ramus; the braincase; the suspensorium; and the parietal bone. The modular pattern of the skull is considered evidence for a flexible framework (this hypothesis conflicts with inferences based on Finite Element Analysis (FEA) modeling;
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" box="[704,891,1908,1934]" journalOrPublisher="Anatomical Record" pageId="22" pageNumber="23" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[704,891,1908,1934]" italics="true" pageId="22" pageNumber="23">Cost et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
); in terms of growth, the suspensorium, parietal, and braincase are the modules that change the least (less than 10 changes per growth stage), which is consistent with their stable keystone-like function against which the relatively flexible palate and facial skeleton can passively move and the lower jaws freely rotate.
|
||
</paragraph>
|
||
<paragraph blockId="23.[498,1542,234,938]" pageId="23" pageNumber="24">
|
||
The greatest change across modules (more than 10) occurs in the 5th and 6th growth stages; this corresponds to the transition from small to large juveniles (
|
||
<figureCitation box="[1338,1417,433,460]" captionStart="Figure 11" captionStartId="22.[524,589,695,717]" captionTargetBox="[502,1535,236,665]" captionTargetId="figure-171@22.[497,1130,233,669]" captionTargetPageId="22" captionText="Figure 11 The frequency distribution of changes to the craniomandibular functional modules (sensu Werneburg et al., 2019) in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. The onset of the changes to the skull roof, snout, mandibular ramus, and suspensorium modules occur early in growth, whereas the onset of changes to the parietal and braincase occur in adulthood. Changes continue throughout growth in all domains, aside from those to the parietal that cease at growth stage 14. Full-size DOI: 10.7717/peerj.9192/fig-11" figureDoi="http://doi.org/10.5281/zenodo.5824994" httpUri="https://zenodo.org/record/5824994/files/figure.png" pageId="23" pageNumber="24">Fig. 11</figureCitation>
|
||
). The transition from the long and low skulls of juveniles to the tall skulls of subadults imposes 52 changes, where the snout dorsum module is changed from the delicate and thin morphotype of juveniles to the greatly expanded and inflated form of subadults.
|
||
</paragraph>
|
||
<paragraph blockId="23.[498,1542,234,938]" pageId="23" pageNumber="24">Changes to the skull roof and mandible are seen throughout ontogeny; changes to the snout and supensorium occur later, from the 5th growth stage onwards. Other modules do not change until young adulthood, including the parietal and braincase; changes to the parietal are limited to young adults and adults, whereas changes to the braincase continue to the last growth stage. These differences in the timing of change to modules is consistent with the hypothesis of modularity, otherwise they would change together if they were integrated. During the interval of greatest change, in the transition from juveniles to subadults (growth stages 5 and 6), the modules that change the most are the skull roof, snout, and mandible.</paragraph>
|
||
<paragraph blockId="23.[498,1542,977,1526]" box="[498,1017,977,1005]" pageId="23" pageNumber="24">
|
||
<heading bold="true" box="[498,1017,977,1005]" fontSize="11" level="3" pageId="23" pageNumber="24" reason="6">
|
||
<emphasis bold="true" box="[498,1017,977,1005]" italics="true" pageId="23" pageNumber="24">Ontogram mapped onto growth curve</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="23.[498,1542,977,1526]" pageId="23" pageNumber="24">
|
||
Mapping the ontogram onto a previously published growth curve for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1295,1361,1021,1046]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1295,1361,1021,1046]" italics="true" pageId="23" pageNumber="24">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" journalOrPublisher="Nature" pageId="23" pageNumber="24" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="23" pageNumber="24">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) permits refinement of the diagnosis of the higher-level growth categories of
|
||
<bibRefCitation author="Carr TD" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="23" pageNumber="24" pagination="497 - 520" part="19" refId="ref52638" refString="Carr TD. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria: Coelurosauria). Journal of Vertebrate Paleontology 19 (3): 497 - 520 DOI 10.1080 / 02724634.1999.10011161." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria: Coelurosauria)" type="journal article" year="1999">
|
||
<emphasis italics="true" pageId="23" pageNumber="24">Carr (1999)</emphasis>
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[634,880,1100,1126]" journalOrPublisher="Nature" pageId="23" pageNumber="24" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[634,880,1100,1126]" italics="true" pageId="23" pageNumber="24">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
, namely juvenile, subadult, young adult, adult, and senescent adult (
|
||
<figureCitation box="[696,775,1140,1166]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="23" pageNumber="24">Fig. 12</figureCitation>
|
||
). This approach provides a framework for comparison with previous studies on functional morphology in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1046,1115,1181,1206]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1046,1115,1181,1206]" italics="true" pageId="23" pageNumber="24">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and the evolution of its ontogeny. This template serves as a heuristic device and is not intended to estimate growth rates; for example, the position of
|
||
<materialsCitation box="[791,974,1259,1286]" collectionCode="BMRP" pageId="23" pageNumber="24" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
on the steepest part of the growth curve conflicts with the histological evidence that the individual was at a moderate growth rate before death (
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[663,928,1339,1366]" journalOrPublisher="Science Advances" pageId="23" pageNumber="24" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[663,928,1339,1366]" italics="true" pageId="23" pageNumber="24">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
). At best, the curve is an approximation of the true growth rate. The chronological age of
|
||
<materialsCitation box="[950,1133,1379,1406]" collectionCode="BMRP" pageId="23" pageNumber="24" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
is based on the revised age of
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[498,775,1419,1445]" journalOrPublisher="Science Advances" pageId="23" pageNumber="24" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[498,775,1419,1445]" italics="true" pageId="23" pageNumber="24">Woodward et al. (2020)</emphasis>
|
||
</bibRefCitation>
|
||
; the age of
|
||
<materialsCitation box="[915,1074,1419,1446]" collectionCode="LACM" pageId="23" pageNumber="24" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
(14 years;
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1204,1439,1419,1445]" journalOrPublisher="Nature" pageId="23" pageNumber="24" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1204,1439,1419,1445]" italics="true" pageId="23" pageNumber="24">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) is considered here to be an underestimate since it conflicts with that of the less mature
|
||
<materialsCitation collectionCode="BMRP" pageId="23" pageNumber="24" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
(15 years;
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[725,990,1499,1525]" journalOrPublisher="Science Advances" pageId="23" pageNumber="24" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[725,990,1499,1525]" italics="true" pageId="23" pageNumber="24">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="23.[498,1541,1570,1921]" box="[498,776,1570,1600]" pageId="23" pageNumber="24">
|
||
<heading bold="true" box="[498,776,1570,1600]" fontSize="12" level="2" pageId="23" pageNumber="24" reason="0">
|
||
<emphasis bold="true" box="[498,776,1570,1600]" pageId="23" pageNumber="24">Growth categories</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="23.[498,1541,1570,1921]" lastBlockId="25.[498,1465,967,1033]" lastPageId="25" lastPageNumber="26" pageId="23" pageNumber="24">
|
||
Alignment between the growth series, chronological age, cortical histology, number of growth changes, growth rate, and, to a lesser degree, size and mass, were used to define five growth categories, once the ontogram was mapped onto the growth curve of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" journalOrPublisher="Nature" pageId="23" pageNumber="24" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="23" pageNumber="24">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
, a procedure that was constrained by seven histologically aged specimens (
|
||
<figureCitation box="[508,588,1775,1801]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="23" pageNumber="24">Fig. 12</figureCitation>
|
||
). Although mass is a relevant variable in these comparisons, given the variety of methods used by different workers, the published mass estimates conflict with each other (
|
||
<tableCitation box="[579,666,1855,1882]" captionStart="Table 5" captionStartId="25.[117,172,248,269]" captionTargetBox="[108,1524,407,885]" captionText="Table 5 Summary of mass estimates for Tyrannosaurus rex. Summary of published mass estimates (kg) for individual specimens of Tyrannosaurus rex with rank order in each column in parentheses; sum rank order given in right-hand column. The rank of AMNH FARB 5027 could not be identified given its presence in only one analysis and the great difference in estimates between the analyses. Column numbers: 1, Erickson et al. (2004); 2, Henderson & Snively (2004); 3, Bates et al. (2009); 4, Hutchinson et al. (2011); 5, Campione et al. (2014); 6, Snively et al. (2019); 7, Persons, Currie & Erickson (2019)." httpUri="http://table.plazi.org/id/DF5C84E2FFDDFFB1FF93FF703676FE3F" pageId="23" pageNumber="24" tableUuid="DF5C84E2FFDDFFB1FF93FF703676FE3F">Table 5</tableCitation>
|
||
); for that reason, the decision for each mention of mass as a relevant diagnostic character is given below. Also, although the masses are given to the nearest kilogram or, in four instances, to the tenth of a kilogram, it is not intended to give the impression of precision, rather the exact estimates are here simply reproduced unaltered from the original sources. To reflect the natural imprecision, mass estimates given in the next section are rounded up in the discussion of diagnostic features of growth categories.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824996" ID-Zenodo-Dep="5824996" httpUri="https://zenodo.org/record/5824996/files/figure.png" pageId="24" pageNumber="25" startId="24.[524,589,1020,1042]" targetBox="[485,1542,220,1000]" targetPageId="24">
|
||
<paragraph blockId="24.[524,1516,1020,1773]" pageId="24" pageNumber="25">
|
||
<emphasis bold="true" pageId="24" pageNumber="25">
|
||
Figure 12
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[630,815,1021,1042]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[630,815,1021,1042]" italics="true" pageId="24" pageNumber="25">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
ontogram, chronological age, and mass mapped onto the growth curve of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[549,756,1049,1071]" journalOrPublisher="Nature" pageId="24" pageNumber="25" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis bold="true" box="[549,756,1049,1071]" italics="true" pageId="24" pageNumber="25">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
.
|
||
</emphasis>
|
||
Ranges of growth categories are indicated across the top. Each circle represents an individual specimen; the vertical columns of circles indicate multiple specimens in a single growth stage; the horizontal position of the white circles does not imply a specific chronological age. For ease of interpretation, and to accommodate missing mass estimates, the position of the circles do not correspond with the scale of the
|
||
<emphasis box="[721,731,1167,1188]" italics="true" pageId="24" pageNumber="25">y</emphasis>
|
||
-axis. The gray circles indicate histologically aged specimens that are connected to the
|
||
<emphasis box="[562,573,1197,1218]" italics="true" pageId="24" pageNumber="25">x</emphasis>
|
||
-axis by dashed lines for ease of interpretation. The star represents the ~3,000 kg threshold that separates
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[616,670,1226,1247]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[616,670,1226,1247]" italics="true" pageId="24" pageNumber="25">T. rex</emphasis>
|
||
</taxonomicName>
|
||
from its closest, but smaller, relatives. Alternating gray and white bars in the background block out the growth categories and their blurred edges reflect the imprecision of their boundaries. The height of each black vertical bar corresponds to the number of synontomorphies in each growth stage, which ranges from 1 to 90 characters. Asterisk indicates the type specimen of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1373,1432,1314,1335]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1373,1432,1314,1335]" italics="true" pageId="24" pageNumber="25">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Female symbols indicate BMRP 2006.6.4 and
|
||
<materialsCitation box="[879,984,1342,1364]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, the only unambiguous female specimens in the data set. Skull illustrations are to scale, with that of the adult set to 1.3 m. From left to right the skulls are:
|
||
<materialsCitation collectionCode="CMNH" pageId="24" pageNumber="25" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
,
|
||
<materialsCitation box="[582,733,1401,1422]" collectionCode="BMRP" pageId="24" pageNumber="25" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
, and
|
||
<materialsCitation box="[788,982,1400,1422]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="24" pageNumber="25" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
. The large juvenile
|
||
<materialsCitation box="[1173,1322,1401,1422]" collectionCode="BMRP" pageId="24" pageNumber="25" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
is in dorsal view to show the early occurrence of the narrow snout and wide temporal region that characterizes
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1399,1453,1430,1451]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1399,1453,1430,1451]" italics="true" pageId="24" pageNumber="25">T. rex</emphasis>
|
||
</taxonomicName>
|
||
to the exclusion of other tyrannosaurids, which is ontogenetically incongruent with its late-occurring (i.e., autapomorphic) phylogenetic homolog; see text for discussion. Major life history events are indicated, including the onset of sexual maturity and somatic maturity, and the earliest occurrence of histologic adulthood. Suspensorium in
|
||
<materialsCitation box="[876,1003,1546,1568]" collectionCode="CMNH" pageId="24" pageNumber="25" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
is reconstructed after
|
||
<materialsCitation box="[1222,1374,1547,1568]" collectionCode="BMRP" pageId="24" pageNumber="25" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
. EFS, External Fundamental System. Specimens: 1,
|
||
<materialsCitation box="[865,995,1576,1597]" collectionCode="LACM" pageId="24" pageNumber="25" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
; 2,
|
||
<materialsCitation box="[1025,1214,1576,1598]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47787" pageId="24" pageNumber="25" specimenCode="AMNH FARB 5050">AMNH FARB 5050</materialsCitation>
|
||
; 3,
|
||
<materialsCitation box="[1243,1357,1576,1597]" collectionCode="DDM" pageId="24" pageNumber="25" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
; 4,
|
||
<materialsCitation box="[1386,1511,1576,1598]" collectionCode="CMNH" pageId="24" pageNumber="25" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
; 5, BMRP 2002.4.1; 6,
|
||
<materialsCitation box="[752,874,1605,1626]" collectionCode="RSM" pageId="24" pageNumber="25" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
; 7, BMRP 2006.4.4; 8,
|
||
<materialsCitation box="[1118,1254,1605,1627]" collectionCode="LACM" pageId="24" pageNumber="25" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
; 9,
|
||
<materialsCitation box="[1298,1410,1605,1627]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
; 10, TMP 1981.006.0001; 11,
|
||
<materialsCitation box="[707,849,1634,1656]" collectionCode="LACM" pageId="24" pageNumber="25" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
; 12,
|
||
<materialsCitation box="[895,1087,1634,1656]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="24" pageNumber="25" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
; 13,
|
||
<materialsCitation box="[1131,1273,1634,1656]" collectionCode="UWBM" pageId="24" pageNumber="25" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
; 14, TMP 1981.012.0001; 15,
|
||
<materialsCitation box="[558,674,1663,1685]" collectionCode="RSM" pageId="24" pageNumber="25" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
; 16,
|
||
<materialsCitation box="[718,843,1664,1685]" collectionCode="SDSM" pageId="24" pageNumber="25" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
; 17,
|
||
<materialsCitation box="[887,1077,1663,1685]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="24" pageNumber="25" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
; 18,
|
||
<materialsCitation box="[1120,1311,1663,1685]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47763" pageId="24" pageNumber="25" specimenCode="AMNH FARB 5029">AMNH FARB 5029</materialsCitation>
|
||
; 19,
|
||
<materialsCitation box="[1353,1512,1664,1685]" collectionCode="NHMUK" pageId="24" pageNumber="25" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
; 20,
|
||
<materialsCitation box="[558,730,1693,1715]" collectionCode="NMMNH" pageId="24" pageNumber="25" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
; 21,
|
||
<materialsCitation box="[775,882,1693,1714]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
; 22,
|
||
<materialsCitation box="[927,1024,1693,1714]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
; 23,
|
||
<materialsCitation box="[1069,1165,1693,1715]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
; 24,
|
||
<materialsCitation box="[1210,1341,1693,1714]" collectionCode="LACM" pageId="24" pageNumber="25" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
; 25,
|
||
<materialsCitation box="[1386,1477,1693,1714]" collectionCode="CM" pageId="24" pageNumber="25" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
; 26,
|
||
<materialsCitation box="[524,668,1722,1743]" collectionCode="UCMP" pageId="24" pageNumber="25" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
; 27,
|
||
<materialsCitation box="[709,804,1722,1743]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
; 28,
|
||
<materialsCitation box="[846,985,1722,1743]" collectionCode="UMNH" pageId="24" pageNumber="25" specimenCode="UMNH 11000">UMNH 11000</materialsCitation>
|
||
; 29,
|
||
<materialsCitation box="[1027,1134,1722,1743]" collectionCode="MOR" pageId="24" pageNumber="25" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
; 30,
|
||
<materialsCitation box="[1175,1295,1722,1743]" collectionCode="UWGM" pageId="24" pageNumber="25" specimenCode="UWGM 181">UWGM 181</materialsCitation>
|
||
; 31,
|
||
<materialsCitation box="[1337,1488,1722,1743]" collectionCode="FMNH" pageId="24" pageNumber="25" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="24.[524,1516,1020,1773]" box="[1099,1515,1751,1773]" pageId="24" pageNumber="25">Full-size DOI: 10.7717/peerj.9192/fig-12</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFDDFFB1FF93FF703676FE3F" httpUri="http://table.plazi.org/id/DF5C84E2FFDDFFB1FF93FF703676FE3F" pageId="25" pageNumber="26" startId="25.[117,172,248,269]" targetBox="[108,1524,407,885]" targetIsTable="true" targetPageId="25">
|
||
<paragraph blockId="25.[117,1516,248,387]" pageId="25" pageNumber="26">
|
||
<emphasis bold="true" box="[117,741,248,270]" pageId="25" pageNumber="26">
|
||
Table 5 Summary of mass estimates for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[545,735,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[545,735,249,270]" italics="true" pageId="25" pageNumber="26">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of published mass estimates (kg) for individual specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis italics="true" pageId="25" pageNumber="26">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
with rank order in each column in parentheses; sum rank order given in right-hand column. The rank of
|
||
<materialsCitation box="[1268,1457,277,299]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="25" pageNumber="26" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
could not be identified given its presence in only one analysis and the great difference in estimates between the analyses. Column numbers: 1,
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" journalOrPublisher="Nature" pageId="25" pageNumber="26" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="25" pageNumber="26">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
; 2,
|
||
<bibRefCitation author="Henderson DM & Snively E." box="[211,476,336,357]" journalOrPublisher="Proceedings of the Royal Society B" pageId="25" pageNumber="26" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[211,476,336,357]" italics="true" pageId="25" pageNumber="26">Henderson & Snively (2004)</emphasis>
|
||
</bibRefCitation>
|
||
; 3,
|
||
<bibRefCitation author="Bates KT & Manning PL & Hodgetts D & Sellers WI" box="[509,678,336,358]" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" pagination="e 4532" part="4" refId="ref52064" refString="Bates KT, Manning PL, Hodgetts D, Sellers WI. 2009. Estimating mass properties of dinosaurs using later imaging and 3 D computer modeling. PLOS ONE 4 (2): e 4532 DOI 10.1371 / journal. pone. 0004532." title="Estimating mass properties of dinosaurs using later imaging and 3 D computer modeling" type="journal article" year="2009">
|
||
<emphasis box="[509,678,336,358]" italics="true" pageId="25" pageNumber="26">Bates et al. (2009)</emphasis>
|
||
</bibRefCitation>
|
||
; 4,
|
||
<bibRefCitation author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[711,939,336,358]" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" pagination="e 97055" part="9" refId="ref54676" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLOS ONE 9 (5): e 97055 DOI 10.1371 / journal. pone. 0097055." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">
|
||
<emphasis box="[711,939,336,358]" italics="true" pageId="25" pageNumber="26">Hutchinson et al. (2011)</emphasis>
|
||
</bibRefCitation>
|
||
; 5,
|
||
<bibRefCitation author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[972,1187,336,358]" journalOrPublisher="Methods in Ecology and Evolution" pageId="25" pageNumber="26" pagination="913 - 923" part="5" refId="ref52505" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods in Ecology and Evolution 5 (9): 913 - 923 DOI 10.1111 / 2041 - 210 X. 12226." title="Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">
|
||
<emphasis box="[972,1187,336,358]" italics="true" pageId="25" pageNumber="26">Campione et al. (2014)</emphasis>
|
||
</bibRefCitation>
|
||
; 6,
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[1221,1405,336,358]" journalOrPublisher="PeerJ" pageId="25" pageNumber="26" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[1221,1405,336,358]" italics="true" pageId="25" pageNumber="26">Snively et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
; 7,
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" journalOrPublisher="Anatomical Record" pageId="25" pageNumber="26" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="25" pageNumber="26">Persons, Currie & Erickson (2019)</emphasis>
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="25" pageNumber="26">
|
||
<table box="[108,1524,407,885]" gridcols="9" gridrows="13" pageId="25" pageNumber="26">
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<tr box="[108,1524,407,429]" gridrow="0" pageId="25" pageNumber="26">
|
||
<th box="[108,301,407,429]" gridcol="0" gridrow="0" pageId="25" pageNumber="26">Specimen</th>
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<th box="[365,451,407,429]" gridcol="1" gridrow="0" pageId="25" pageNumber="26">1</th>
|
||
<th box="[515,612,407,429]" gridcol="2" gridrow="0" pageId="25" pageNumber="26">2</th>
|
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<th box="[676,779,407,429]" gridcol="3" gridrow="0" pageId="25" pageNumber="26">3</th>
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||
<th box="[843,941,407,429]" gridcol="4" gridrow="0" pageId="25" pageNumber="26">4</th>
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||
<th box="[1005,1091,407,429]" gridcol="5" gridrow="0" pageId="25" pageNumber="26">5</th>
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||
<th box="[1155,1258,407,429]" gridcol="6" gridrow="0" pageId="25" pageNumber="26">6</th>
|
||
<th box="[1322,1408,407,429]" gridcol="7" gridrow="0" pageId="25" pageNumber="26">7</th>
|
||
<th box="[1472,1524,407,429]" gridcol="8" gridrow="0" pageId="25" pageNumber="26">
|
||
<emphasis bold="true" box="[1472,1524,408,429]" pageId="25" pageNumber="26">Rank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1524,449,471]" gridrow="1" pageId="25" pageNumber="26" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1">
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<th box="[108,301,449,471]" gridcol="0" gridrow="1" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,238,449,470]" collectionCode="LACM" pageId="25" pageNumber="26" specimenCode="LACM 28471">LACM 28471</materialsCitation>
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||
</th>
|
||
<td box="[365,451,449,471]" gridcol="1" gridrow="1" pageId="25" pageNumber="26">29.9 (1)</td>
|
||
<td box="[1472,1524,449,471]" gridcol="8" gridrow="1" pageId="25" pageNumber="26">1</td>
|
||
</tr>
|
||
<tr box="[108,1524,486,509]" gridrow="2" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-5="1" rowspan-7="1">
|
||
<th box="[108,301,486,509]" gridcol="0" gridrow="2" pageId="25" pageNumber="26">
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<materialsCitation box="[108,257,486,508]" collectionCode="BMRP" pageId="25" pageNumber="26" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
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</th>
|
||
<td box="[843,941,486,509]" gridcol="4" gridrow="2" pageId="25" pageNumber="26">954 (1)</td>
|
||
<td box="[1155,1258,486,509]" gridcol="6" gridrow="2" pageId="25" pageNumber="26">660.2 (1)</td>
|
||
<td box="[1472,1524,486,509]" gridcol="8" gridrow="2" pageId="25" pageNumber="26">2</td>
|
||
</tr>
|
||
<tr box="[108,1524,524,546]" gridrow="3" pageId="25" pageNumber="26" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1">
|
||
<th box="[108,301,524,546]" gridcol="0" gridrow="3" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,238,524,546]" collectionCode="LACM" pageId="25" pageNumber="26" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</th>
|
||
<td box="[365,451,524,546]" gridcol="1" gridrow="3" pageId="25" pageNumber="26">1,807 (2)</td>
|
||
<td box="[1472,1524,524,546]" gridcol="8" gridrow="3" pageId="25" pageNumber="26">3</td>
|
||
</tr>
|
||
<tr box="[108,1524,561,584]" gridrow="4" pageId="25" pageNumber="26" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
|
||
<th box="[108,301,561,584]" gridcol="0" gridrow="4" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,301,561,583]" collectionCode="TMP" pageId="25" pageNumber="26" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[365,451,561,584]" gridcol="1" gridrow="4" pageId="25" pageNumber="26">3,230 (3)</td>
|
||
<td box="[1322,1408,561,584]" gridcol="7" gridrow="4" pageId="25" pageNumber="26">4,469 (1)</td>
|
||
<td box="[1472,1524,561,584]" gridcol="8" gridrow="4" pageId="25" pageNumber="26">4</td>
|
||
</tr>
|
||
<tr box="[108,1524,599,621]" gridrow="5" pageId="25" pageNumber="26" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1">
|
||
<th box="[108,301,599,621]" gridcol="0" gridrow="5" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,301,599,621]" collectionCode="TMP" pageId="25" pageNumber="26" specimenCode="TMP 1981.006.0001">TMP 1981.012.0001</materialsCitation>
|
||
</th>
|
||
<td box="[365,451,599,621]" gridcol="1" gridrow="5" pageId="25" pageNumber="26">5,040 (4)</td>
|
||
<td box="[1472,1524,599,621]" gridcol="8" gridrow="5" pageId="25" pageNumber="26">5</td>
|
||
</tr>
|
||
<tr box="[108,1524,637,659]" gridrow="6" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
|
||
<th box="[108,301,637,659]" gridcol="0" gridrow="6" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,202,637,658]" collectionCode="MOR" pageId="25" pageNumber="26" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[1322,1408,637,659]" gridcol="7" gridrow="6" pageId="25" pageNumber="26">5,112 (2)</td>
|
||
<td box="[1472,1524,637,659]" gridcol="8" gridrow="6" pageId="25" pageNumber="26">6</td>
|
||
</tr>
|
||
<tr box="[108,1524,674,696]" gridrow="7" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
|
||
<th box="[108,301,674,696]" gridcol="0" gridrow="7" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,214,674,696]" collectionCode="MOR" pageId="25" pageNumber="26" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[1322,1408,674,696]" gridcol="7" gridrow="7" pageId="25" pageNumber="26">6,100 (3)</td>
|
||
<td box="[1472,1524,674,696]" gridcol="8" gridrow="7" pageId="25" pageNumber="26">7</td>
|
||
</tr>
|
||
<tr box="[108,1524,712,734]" gridrow="8" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-7="1">
|
||
<th box="[108,301,712,734]" gridcol="0" gridrow="8" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,299,712,734]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="25" pageNumber="26" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
</th>
|
||
<td box="[1155,1258,712,734]" gridcol="6" gridrow="8" pageId="25" pageNumber="26">6,986.6 (2)</td>
|
||
<td box="[1472,1524,712,734]" gridcol="8" gridrow="8" pageId="25" pageNumber="26">?</td>
|
||
</tr>
|
||
<tr box="[108,1524,749,772]" gridrow="9" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-6="1">
|
||
<th box="[108,301,749,772]" gridcol="0" gridrow="9" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,202,749,771]" collectionCode="MOR" pageId="25" pageNumber="26" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[676,779,749,772]" gridcol="3" gridrow="9" pageId="25" pageNumber="26">6,071.8 (1)</td>
|
||
<td box="[843,941,749,772]" gridcol="4" gridrow="9" pageId="25" pageNumber="26">8,272 (2)</td>
|
||
<td box="[1005,1091,749,772]" gridcol="5" gridrow="9" pageId="25" pageNumber="26">6,216 (1)</td>
|
||
<td box="[1322,1408,749,772]" gridcol="7" gridrow="9" pageId="25" pageNumber="26">6,264 (4)</td>
|
||
<td box="[1472,1524,749,772]" gridcol="8" gridrow="9" pageId="25" pageNumber="26">8</td>
|
||
</tr>
|
||
<tr box="[108,1524,787,809]" gridrow="10" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-6="1">
|
||
<th box="[108,301,787,809]" gridcol="0" gridrow="10" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,198,787,808]" collectionCode="CM" pageId="25" pageNumber="26" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</th>
|
||
<td box="[843,941,787,809]" gridcol="4" gridrow="10" pageId="25" pageNumber="26">9,081 (3)</td>
|
||
<td box="[1005,1091,787,809]" gridcol="5" gridrow="10" pageId="25" pageNumber="26">6,688 (2)</td>
|
||
<td box="[1322,1408,787,809]" gridcol="7" gridrow="10" pageId="25" pageNumber="26">6,740 (5)</td>
|
||
<td box="[1472,1524,787,809]" gridcol="8" gridrow="10" pageId="25" pageNumber="26">9</td>
|
||
</tr>
|
||
<tr box="[108,1524,825,847]" gridrow="11" pageId="25" pageNumber="26" rowspan-3="1">
|
||
<th box="[108,301,825,847]" gridcol="0" gridrow="11" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,258,825,846]" collectionCode="FMNH" pageId="25" pageNumber="26" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[365,451,825,847]" gridcol="1" gridrow="11" pageId="25" pageNumber="26">5,654 (5)</td>
|
||
<td box="[515,612,825,847]" gridcol="2" gridrow="11" pageId="25" pageNumber="26">10,200 (1)</td>
|
||
<td box="[843,941,825,847]" gridcol="4" gridrow="11" pageId="25" pageNumber="26">13,996 (4)</td>
|
||
<td box="[1005,1091,825,847]" gridcol="5" gridrow="11" pageId="25" pageNumber="26">7,377 (3)</td>
|
||
<td box="[1155,1258,825,847]" gridcol="6" gridrow="11" pageId="25" pageNumber="26">9,130.9 (3)</td>
|
||
<td box="[1322,1408,825,847]" gridcol="7" gridrow="11" pageId="25" pageNumber="26">8,462 (6)</td>
|
||
<td box="[1472,1524,825,847]" gridcol="8" gridrow="11" pageId="25" pageNumber="26">10</td>
|
||
</tr>
|
||
<tr box="[108,1524,862,885]" gridrow="12" pageId="25" pageNumber="26" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-6="1">
|
||
<th box="[108,301,862,885]" gridcol="0" gridrow="12" pageId="25" pageNumber="26">
|
||
<materialsCitation box="[108,224,862,884]" collectionCode="RSM" pageId="25" pageNumber="26" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[1005,1091,862,885]" gridcol="5" gridrow="12" pageId="25" pageNumber="26">8,004 (4)</td>
|
||
<td box="[1322,1408,862,885]" gridcol="7" gridrow="12" pageId="25" pageNumber="26">8,870 (7)</td>
|
||
<td box="[1472,1524,862,885]" gridcol="8" gridrow="12" pageId="25" pageNumber="26">11</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="25.[498,1542,1078,1908]" box="[498,642,1078,1108]" pageId="25" pageNumber="26">
|
||
<heading bold="true" box="[498,642,1078,1108]" fontSize="12" level="2" pageId="25" pageNumber="26" reason="0">
|
||
<emphasis bold="true" box="[498,642,1078,1108]" pageId="25" pageNumber="26">Juveniles</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="25.[498,1542,1078,1908]" pageId="25" pageNumber="26">
|
||
Juveniles (=adolescent category of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[897,1124,1124,1150]" journalOrPublisher="Nature" pageId="25" pageNumber="26" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[897,1124,1124,1150]" italics="true" pageId="25" pageNumber="26">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) have a skull length less than 80 cm, long and low skulls with a length to height ratio of 3.0 or greater, are equal to or less than 13 years old, and lack an EFS. This growth category sees 58 unambiguously optimized growth changes, of which 24 have a CI of 1.0 (
|
||
<tableCitation box="[1052,1140,1243,1269]" captionStart="Table 6" captionStartId="26.[525,580,248,269]" captionTargetBox="[516,1524,378,1306]" captionText="Table 6 Diagnostic characters of the juvenile growth category of Tyrannosaurus rex. These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1)." httpUri="http://table.plazi.org/id/DF5C84E2FFDEFFB2FDEBFF70322CFEDA" pageId="25" pageNumber="26" tableUuid="DF5C84E2FFDEFFB2FDEBFF70322CFEDA">Table 6</tableCitation>
|
||
). In the skull, most changes are seen in the snout module (sensu
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[889,1155,1283,1309]" journalOrPublisher="Scientific Reports" pageId="25" pageNumber="26" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis box="[889,1155,1283,1309]" italics="true" pageId="25" pageNumber="26">Werneburg et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
). This category corresponds to growth stages 1
|
||
<emphasis box="[677,692,1324,1349]" italics="true" pageId="25" pageNumber="26">–</emphasis>
|
||
5 of the ontogram (
|
||
<figureCitation box="[918,982,1323,1349]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="25" pageNumber="26">Fig. 2</figureCitation>
|
||
) and to the initial lag phase of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1385,1452,1324,1349]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="25" pageNumber="26" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1385,1452,1324,1349]" italics="true" pageId="25" pageNumber="26">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth curve of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[603,840,1363,1389]" journalOrPublisher="Nature" pageId="25" pageNumber="26" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[603,840,1363,1389]" italics="true" pageId="25" pageNumber="26">Erickson et al. (2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[855,935,1363,1389]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="25" pageNumber="26">Fig. 12</figureCitation>
|
||
). The juvenile stage extends from hatching to the 13th year of life, before the long and low skull proportions are lost at the subadult category.
|
||
</paragraph>
|
||
<paragraph blockId="25.[498,1542,1078,1908]" pageId="25" pageNumber="26">
|
||
A single mass estimate of 39.9 kg has been published for the small juvenile
|
||
<materialsCitation collectionCode="LACM" pageId="25" pageNumber="26" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
by
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[614,860,1482,1509]" journalOrPublisher="Nature" pageId="25" pageNumber="26" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[614,860,1482,1509]" italics="true" pageId="25" pageNumber="26">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
. Two mass estimates have been published for the large juvenile BMRP 2002.4.1: a range of 639
|
||
<emphasis box="[965,980,1523,1548]" italics="true" pageId="25" pageNumber="26">–</emphasis>
|
||
1,269 kg, and a mean of 954 kg was published by
|
||
<bibRefCitation author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[535,817,1562,1588]" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" pagination="e 97055" part="9" refId="ref54676" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLOS ONE 9 (5): e 97055 DOI 10.1371 / journal. pone. 0097055." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">
|
||
<emphasis box="[535,817,1562,1588]" italics="true" pageId="25" pageNumber="26">Hutchinson et al. (2011)</emphasis>
|
||
</bibRefCitation>
|
||
and an estimate of 660.2 kg was published by
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" journalOrPublisher="PeerJ" pageId="25" pageNumber="26" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[1372,1519,1562,1588]" italics="true" pageId="25" pageNumber="26">Snively et al.</emphasis>
|
||
<emphasis box="[498,572,1602,1628]" italics="true" pageId="25" pageNumber="26">(2019)</emphasis>
|
||
</bibRefCitation>
|
||
. The maximum estimate is used here as a point of reference; therefore, juveniles are here considered to have a mass equal to or less than ~1,300 kg.
|
||
</paragraph>
|
||
<paragraph blockId="25.[498,1542,1078,1908]" lastBlockId="26.[498,1542,1389,1496]" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26">
|
||
The juvenile growth stage is diagnosed by the possession of a dorsoventrally shallow skull and jaws, the increase in height of which defines the subadult stage. To precisely reflect the primary changes that occur even during this initial phase of life, this growth category is divided into two categories in the following diagnosis, namely small juveniles and large juveniles. A total of five growth changes occur in the small juvenile growth stage, which affect the skull roof and mandible modules of
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[1264,1535,1881,1907]" journalOrPublisher="Scientific Reports" pageId="25" pageNumber="26" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis box="[1264,1535,1881,1907]" italics="true" pageId="25" pageNumber="26">Werneburg et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
. In contrast, 53 changes occur in the large juvenile growth stage that are more widespread across the skull, including modules of the skull roof, snout, suspensorium, and mandible. The large juvenile category marks the earliest onset of sexual maturity (see below).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFDEFFB2FDEBFF70322CFEDA" httpUri="http://table.plazi.org/id/DF5C84E2FFDEFFB2FDEBFF70322CFEDA" pageId="26" pageNumber="27" startId="26.[525,580,248,269]" targetBox="[516,1524,378,1306]" targetIsTable="true" targetPageId="26">
|
||
<paragraph blockId="26.[525,1515,248,358]" pageId="26" pageNumber="27">
|
||
<emphasis bold="true" box="[525,1352,248,270]" pageId="26" pageNumber="27">
|
||
Table 6 Diagnostic characters of the juvenile growth category of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1162,1347,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1162,1347,249,270]" italics="true" pageId="26" pageNumber="27">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="26" pageNumber="27">
|
||
<table box="[516,1524,378,1306]" gridcols="2" gridrows="25" pageId="26" pageNumber="27">
|
||
<tr box="[516,1524,378,400]" gridrow="0" pageId="26" pageNumber="27">
|
||
<th box="[516,1205,378,400]" gridcol="0" gridrow="0" pageId="26" pageNumber="27">Character</th>
|
||
<th box="[1372,1524,378,400]" gridcol="1" gridrow="0" pageId="26" pageNumber="27">
|
||
<emphasis bold="true" box="[1372,1524,378,400]" pageId="26" pageNumber="27">Growth change</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,419,442]" gridrow="1" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,419,442]" gridcol="0" gridrow="1" pageId="26" pageNumber="27">4. Orbital fenestra, shape (PHYLO 7)</td>
|
||
<td box="[1372,1524,419,442]" gridcol="1" gridrow="1" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,457,479]" gridrow="2" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,457,479]" gridcol="0" gridrow="2" pageId="26" pageNumber="27">27. Maxilla, maxillary flange, size (PHYLO 47)</td>
|
||
<td box="[1372,1524,457,479]" gridcol="1" gridrow="2" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,494,517]" gridrow="3" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,494,517]" gridcol="0" gridrow="3" pageId="26" pageNumber="27">57. Jugal, postorbital process, orientation (PHYLO 101)</td>
|
||
<td box="[1372,1524,494,517]" gridcol="1" gridrow="3" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,532,554]" gridrow="4" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,532,554]" gridcol="0" gridrow="4" pageId="26" pageNumber="27">66. Postorbital, jugal, ramus, subocular process, presence (PHYLO 116)</td>
|
||
<td box="[1372,1524,532,554]" gridcol="1" gridrow="4" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,570,592]" gridrow="5" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,570,592]" gridcol="0" gridrow="5" pageId="26" pageNumber="27">89. Frontal, postorbital buttress, distinctiveness (PHYLO 159)</td>
|
||
<td box="[1372,1524,570,592]" gridcol="1" gridrow="5" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,607,630]" gridrow="6" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,607,630]" gridcol="0" gridrow="6" pageId="26" pageNumber="27">123. Dentary, Meckelian groove, depth (PHYLO 236)</td>
|
||
<td box="[1372,1524,607,630]" gridcol="1" gridrow="6" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,645,667]" gridrow="7" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,645,667]" gridcol="0" gridrow="7" pageId="26" pageNumber="27">215. Bony naris, dorsal margin</td>
|
||
<td box="[1372,1524,645,667]" gridcol="1" gridrow="7" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,683,705]" gridrow="8" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,683,705]" gridcol="0" gridrow="8" pageId="26" pageNumber="27">226. Internal antorbital fenestra, jugal contribution</td>
|
||
<td box="[1372,1524,683,705]" gridcol="1" gridrow="8" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,720,742]" gridrow="9" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,720,742]" gridcol="0" gridrow="9" pageId="26" pageNumber="27">238. Postorbital bar, jugopostorbital suture</td>
|
||
<td box="[1372,1524,720,742]" gridcol="1" gridrow="9" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,758,780]" gridrow="10" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,758,780]" gridcol="0" gridrow="10" pageId="26" pageNumber="27">239. Postorbital bar, orientation</td>
|
||
<td box="[1372,1524,758,780]" gridcol="1" gridrow="10" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,795,817]" gridrow="11" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,795,817]" gridcol="0" gridrow="11" pageId="26" pageNumber="27">240. Postorbital bar, plane change</td>
|
||
<td box="[1372,1524,795,817]" gridcol="1" gridrow="11" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,833,855]" gridrow="12" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,833,855]" gridcol="0" gridrow="12" pageId="26" pageNumber="27">254. Premaxilla, body, rostral margin, angulation position</td>
|
||
<td box="[1372,1524,833,855]" gridcol="1" gridrow="12" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,871,893]" gridrow="13" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,871,893]" gridcol="0" gridrow="13" pageId="26" pageNumber="27">293. Nasal, cross-section of rostral third, shape</td>
|
||
<td box="[1372,1524,871,893]" gridcol="1" gridrow="13" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,908,930]" gridrow="14" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,908,930]" gridcol="0" gridrow="14" pageId="26" pageNumber="27">296. Nasal, midline bumps, number</td>
|
||
<td box="[1372,1524,908,930]" gridcol="1" gridrow="14" pageId="26" pageNumber="27">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,946,968]" gridrow="15" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,946,968]" gridcol="0" gridrow="15" pageId="26" pageNumber="27">299. Nasal, dorsal foramina, number rows</td>
|
||
<td box="[1372,1524,946,968]" gridcol="1" gridrow="15" pageId="26" pageNumber="27">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,983,1005]" gridrow="16" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,983,1005]" gridcol="0" gridrow="16" pageId="26" pageNumber="27">315. Nasal, frontal ramus, dorsum, cross section</td>
|
||
<td box="[1372,1524,983,1005]" gridcol="1" gridrow="16" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1021,1043]" gridrow="17" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1021,1043]" gridcol="0" gridrow="17" pageId="26" pageNumber="27">358. Maxilla, subcutaneous surface, orientation to ventral margin</td>
|
||
<td box="[1372,1524,1021,1043]" gridcol="1" gridrow="17" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1058,1081]" gridrow="18" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1058,1081]" gridcol="0" gridrow="18" pageId="26" pageNumber="27">415. Maxilla, palatal process, extent of bony choana</td>
|
||
<td box="[1372,1524,1058,1081]" gridcol="1" gridrow="18" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1096,1118]" gridrow="19" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1096,1118]" gridcol="0" gridrow="19" pageId="26" pageNumber="27">556. Jugal, body, degree of texture</td>
|
||
<td box="[1372,1524,1096,1118]" gridcol="1" gridrow="19" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1134,1156]" gridrow="20" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1134,1156]" gridcol="0" gridrow="20" pageId="26" pageNumber="27">621. Postorbital, body, medial rugosities</td>
|
||
<td box="[1372,1524,1134,1156]" gridcol="1" gridrow="20" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1171,1193]" gridrow="21" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1171,1193]" gridcol="0" gridrow="21" pageId="26" pageNumber="27">1069. Dentary, bone, shallowest point</td>
|
||
<td box="[1372,1524,1171,1193]" gridcol="1" gridrow="21" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1209,1231]" gridrow="22" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1209,1231]" gridcol="0" gridrow="22" pageId="26" pageNumber="27">1170. Prearticular, caudal ramus, dorsal & ventral margins, orientation</td>
|
||
<td box="[1372,1524,1209,1231]" gridcol="1" gridrow="22" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1247,1269]" gridrow="23" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1247,1269]" gridcol="0" gridrow="23" pageId="26" pageNumber="27">1173. Prearticular, rostral ramus, form</td>
|
||
<td box="[1372,1524,1247,1269]" gridcol="1" gridrow="23" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1284,1306]" gridrow="24" pageId="26" pageNumber="27">
|
||
<td box="[516,1205,1284,1306]" gridcol="0" gridrow="24" pageId="26" pageNumber="27">1176. Dentition, frst maxllary tooth, form</td>
|
||
<td box="[1372,1524,1284,1306]" gridcol="1" gridrow="24" pageId="26" pageNumber="27">0 ==> 1</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="26.[498,1542,1532,1686]" box="[498,726,1532,1562]" pageId="26" pageNumber="27">
|
||
<heading bold="true" box="[498,726,1532,1562]" fontSize="12" level="2" pageId="26" pageNumber="27" reason="0">
|
||
<emphasis bold="true" box="[498,726,1532,1562]" pageId="26" pageNumber="27">Small juveniles</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="26.[498,1542,1532,1686]" box="[498,695,1576,1604]" pageId="26" pageNumber="27">
|
||
<heading bold="true" box="[498,695,1576,1604]" fontSize="11" level="3" pageId="26" pageNumber="27" reason="6">
|
||
<emphasis bold="true" box="[498,695,1576,1604]" italics="true" pageId="26" pageNumber="27">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="26.[498,1542,1532,1686]" pageId="26" pageNumber="27">In the fourth growth stage, there is a clear distinction between the postorbital buttress and the caudal shelf of the frontal bone.</paragraph>
|
||
<paragraph blockId="26.[498,1542,1725,1875]" box="[498,741,1725,1753]" pageId="26" pageNumber="27">
|
||
<heading bold="true" box="[498,741,1725,1753]" fontSize="11" level="3" pageId="26" pageNumber="27" reason="6">
|
||
<emphasis bold="true" box="[498,741,1725,1753]" italics="true" pageId="26" pageNumber="27">Neurovasculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="26.[498,1542,1725,1875]" pageId="26" pageNumber="27">In the second growth stage, the alveolar row of foramina that penetrates the lateral surface of the dentary acquires a sulcus that unites all of the openings. In medial view, the Meckelian sulcus becomes a deeply inset groove.</paragraph>
|
||
<paragraph blockId="27.[498,1542,233,383]" box="[498,787,233,261]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,787,233,261]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="6">
|
||
<emphasis bold="true" box="[498,787,233,261]" italics="true" pageId="27" pageNumber="28">Dorsotemporal fossa</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,233,383]" pageId="27" pageNumber="28">In the fourth growth stage, the sagittal crest of the frontal becomes tall and the fossa extends rostrodorsally at a steep angle relative to the level forehead (subcutaneous surface ahead of the fossa).</paragraph>
|
||
<paragraph blockId="27.[498,1542,420,1331]" box="[498,728,420,450]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,728,420,450]" fontSize="12" level="2" pageId="27" pageNumber="28" reason="0">
|
||
<emphasis bold="true" box="[498,728,420,450]" pageId="27" pageNumber="28">Large juveniles</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,420,1331]" box="[498,655,464,492]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,655,464,492]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="6">
|
||
<emphasis bold="true" box="[498,655,464,492]" italics="true" pageId="27" pageNumber="28">Skull frame</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,420,1331]" pageId="27" pageNumber="28">In the fifth growth stage, several correlates of an increase in skull height are seen, including the dorsal position of the angulation in the rostral margin of the premaxilla, the rostrodorsal orientation of the nasal process of the premaxilla, an oval orbital fenestra, vertical orientation of the postorbital bar with the presence of a depression and strut on the lateral surface of the postorbital process of the jugal, the inset and braced ventral end of the jugal process of the postorbital, and presence of the subocular process of the postorbital.</paragraph>
|
||
<paragraph blockId="27.[498,1542,420,1331]" pageId="27" pageNumber="28">
|
||
Some changes in skull architecture almost certainly reflect a response to high bite forces (
|
||
<bibRefCitation author="Henderson DM" box="[508,703,827,853]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="27" pageNumber="28" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[508,703,827,853]" italics="true" pageId="27" pageNumber="28">Henderson, 2002</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[718,1027,827,853]" journalOrPublisher="Biology Letters" pageId="27" pageNumber="28" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[718,1027,827,853]" italics="true" pageId="27" pageNumber="28">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[1042,1097,827,853]" journalOrPublisher="Biology Letters" pageId="27" pageNumber="28" pagination="20180160" part="14" refId="ref52146" refString="Bates KT, Falkingham PL. 2018. Correction to ' estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics '. Biology Letters 14 (4): 20180160 DOI 10.1098 / rsbl. 2018.0160." title="Correction to ' estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2018">
|
||
<emphasis box="[1042,1097,827,853]" italics="true" pageId="27" pageNumber="28">2018</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[1112,1401,827,853]" journalOrPublisher="Scientific Reports" pageId="27" pageNumber="28" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis box="[1112,1401,827,853]" italics="true" pageId="27" pageNumber="28">Gignac & Erickson, 2017</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" journalOrPublisher="Anatomical Record" pageId="27" pageNumber="28" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="27" pageNumber="28">Cost et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
) and their corresponding high loads upon the skull and teeth. These include a convex cross section of the rostral third of the nasal and its frontal ramus, the tall height of the antorbital fossa below the internal antorbital fenestra, the straight rostral margin of the rostroventral ala of the lacrimal, and the concave dorsal margin of the vomer.
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,420,1331]" pageId="27" pageNumber="28">Other changes are almost certainly unrelated to the increase in skull height, including the straight dorsal margin of the bony naris, the presence of a contribution of the jugal to the margin of the internal antorbital fenestra, enhancement of the plane change between the temporal region and the orbitosnout region, the medioventral orientation of the lateral surface of the maxilla to the alveolar margin of the bone, the intermediate position of the bony choana, the rostralward shift of the ventral margin of the joint surface for the quadratojugal ramus on the jugal, the straight shape of the dentary in ventral view, and caudalward position of the shallowest point of the dentary.</paragraph>
|
||
<paragraph blockId="27.[498,1542,1369,1519]" box="[498,695,1369,1397]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,695,1369,1397]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="6">
|
||
<emphasis bold="true" box="[498,695,1369,1397]" italics="true" pageId="27" pageNumber="28">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,1369,1519]" pageId="27" pageNumber="28">In the fifth growth stage an alteration for a stable skull frame is seen, namely a distinct maxillary flange that produces a deep slot for the nasal. The plane change between snout and orbitotemporal region is reflected in the sinuous jugopostorbital suture.</paragraph>
|
||
<paragraph blockId="27.[498,1541,1557,1747]" box="[498,809,1557,1585]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,809,1557,1585]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="6">
|
||
<emphasis bold="true" box="[498,809,1557,1585]" italics="true" pageId="27" pageNumber="28">Subcutaneous surface</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1541,1557,1747]" pageId="27" pageNumber="28">
|
||
In the fifth growth stage, an increase in the relief of the subcutaneous surface of the face is seen, where alveolar skirts are present at the premaxilla, the body of the jugal is rugose, indicating the appearance of armor-like skin (
|
||
<bibRefCitation author="Carr TD & Varricchio DJ & Sedlmayr JC & Roberts EM & Moore JR" box="[1041,1231,1680,1707]" journalOrPublisher="Scientific Reports" pageId="27" pageNumber="28" pagination="44942" part="7" refId="ref52780" refString="Carr TD, Varricchio DJ, Sedlmayr JC, Roberts EM, Moore JR. 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7 (1): 44942 DOI 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">
|
||
<emphasis box="[1041,1231,1680,1707]" italics="true" pageId="27" pageNumber="28">Carr et al., 2017</emphasis>
|
||
</bibRefCitation>
|
||
), and subcutaneous rugosities extend onto the medial surface of the postorbital.
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,1784,1935]" box="[498,828,1784,1812]" pageId="27" pageNumber="28">
|
||
<heading bold="true" box="[498,828,1784,1812]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="6">
|
||
<emphasis bold="true" box="[498,828,1784,1812]" italics="true" pageId="27" pageNumber="28">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="27.[498,1542,1784,1935]" pageId="27" pageNumber="28">In the fifth growth stage, some changes in cephalic ornamentation are independent of the texture of the subcutaneous surface, including the decrease in size of the cornual process of the jugal and the increase in the number of midline bumps on the nasals.</paragraph>
|
||
<paragraph blockId="28.[498,1542,233,622]" box="[498,822,233,261]" pageId="28" pageNumber="29">
|
||
<heading bold="true" box="[498,822,233,261]" fontSize="11" level="3" pageId="28" pageNumber="29" reason="6">
|
||
<emphasis bold="true" box="[498,822,233,261]" italics="true" pageId="28" pageNumber="29">Paranasal pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="28.[498,1542,233,622]" pageId="28" pageNumber="29">In the fifth growth stage, internal inflation of the maxillary sinus is seen externally, shown by the convex rostrodorsal surface of the bone. Some changes to the antorbital fossa might correspond to internal changes of the dentition, including elimination of the rostral end of the crease that defines the ventral margin of the external antorbital fenestra.</paragraph>
|
||
<paragraph blockId="28.[498,1542,233,622]" pageId="28" pageNumber="29">In the maxilla, the rostroventral corner of the external antorbital fossa is indistinct, but it is difficult to determine whether this is caused by a difference in pneumaticity or of the subcutaneous surface; the antorbital fossa is deeply inset relative to the lateral surface of the bone, especially rostrodorsally; and the rostral pneumatic recess of the palatine becomes longer than the caudal recess.</paragraph>
|
||
<paragraph blockId="28.[498,1542,661,771]" box="[498,741,661,689]" pageId="28" pageNumber="29">
|
||
<heading bold="true" box="[498,741,661,689]" fontSize="11" level="3" pageId="28" pageNumber="29" reason="6">
|
||
<emphasis bold="true" box="[498,741,661,689]" italics="true" pageId="28" pageNumber="29">Neurovasculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="28.[498,1542,661,771]" pageId="28" pageNumber="29">In the fifth growth stage, the number of foramina rows in the nasal increases to three and the rostrodorsal foramen of the tract of the subnarial foramen makes its appearance.</paragraph>
|
||
<paragraph blockId="28.[498,1542,809,1198]" box="[498,807,809,837]" pageId="28" pageNumber="29">
|
||
<heading bold="true" box="[498,807,809,837]" fontSize="11" level="3" pageId="28" pageNumber="29" reason="6">
|
||
<emphasis bold="true" box="[498,807,809,837]" italics="true" pageId="28" pageNumber="29">Adductor musculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="28.[498,1542,809,1198]" pageId="28" pageNumber="29">In the fifth growth stage, the insertion for the ventral pterygoid muscle on the surangular and dentary is enhanced, the dorsal margin of the postorbital is everted medially, and in rostral and medial views its dorsal margin is convex.</paragraph>
|
||
<paragraph blockId="28.[498,1542,809,1198]" pageId="28" pageNumber="29">Several bony changes almost certainly reflect an increase in the forcefulness of the adductor musculature, including rostrally converging dorsal and ventral margins of the caudal ramus of the prearticular. Changes in the adductor musculature impose alterations to bones that are not directly related to increased bite force; for example, the rostral ramus of the prearticular is paddle-shaped, which in life medially increased the bony enclosure around the inserted musculature.</paragraph>
|
||
<paragraph blockId="28.[498,1542,1237,1387]" box="[498,700,1237,1265]" pageId="28" pageNumber="29">
|
||
<heading bold="true" box="[498,700,1237,1265]" fontSize="11" level="3" pageId="28" pageNumber="29" reason="6">
|
||
<emphasis bold="true" box="[498,700,1237,1265]" italics="true" pageId="28" pageNumber="29">Joint capsules</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="28.[498,1542,1237,1387]" pageId="28" pageNumber="29">In the fifth growth stage, the horizontal ridge on the ventral quadrate process of the quadratojugal is absent, and the lateral scar below the glenoid of the surangular becomes rugose.</paragraph>
|
||
<paragraph blockId="28.[498,1542,1426,1894]" box="[498,623,1426,1454]" pageId="28" pageNumber="29">
|
||
<heading bold="true" box="[498,623,1426,1454]" fontSize="11" level="3" pageId="28" pageNumber="29" reason="6">
|
||
<emphasis bold="true" box="[498,623,1426,1454]" italics="true" pageId="28" pageNumber="29">Dentition</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="28.[498,1542,1426,1894]" pageId="28" pageNumber="29">
|
||
In the fifth growth stage, the maxillary tooth count increases to 16 and the dentary tooth count increases to 17; and the first maxillary tooth becomes subincisiform. The teeth and antorbital fossa complex are useful points of reference for capturing changes in tooth size. For example, in the fifth growth stage less than six teeth are ahead of the external antorbital fossa, which almost certainly reflects enlargement of the teeth if, and only if, the rostral margin of the fenestra is a stable landmark throughout ontogeny. Geometric morphometric quantification of skull shape in
|
||
<taxonomicName box="[1046,1154,1709,1735]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1046,1154,1709,1735]" italics="true" pageId="28" pageNumber="29">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
shows that the rostral end of the fossa shifts caudally during growth (
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[928,1303,1748,1775]" journalOrPublisher="PeerJ" pageId="28" pageNumber="29" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[928,1303,1748,1775]" italics="true" pageId="28" pageNumber="29">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
). If the same trend occurs in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[615,684,1789,1814]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[615,684,1789,1814]" italics="true" pageId="28" pageNumber="29">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and if there was no increase in the size of teeth then an increase in the number of teeth ahead of the fossa would be seen. Therefore, tooth size increases despite the caudalward shift of the fossa.
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,233,1421]" box="[498,651,233,263]" pageId="29" pageNumber="30">
|
||
<heading bold="true" box="[498,651,233,263]" fontSize="12" level="2" pageId="29" pageNumber="30" reason="0">
|
||
<emphasis bold="true" box="[498,651,233,263]" pageId="29" pageNumber="30">Subadults</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,233,1421]" pageId="29" pageNumber="30">
|
||
Subadults (= juvenile category of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[892,1126,279,305]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[892,1126,279,305]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) have a skull length from 80 cm to 1.1 m, tall skulls with a length to height ratio less than 3.0 (deduced from the straight form of the ventral ramus of the lacrimal in
|
||
<materialsCitation box="[959,1118,358,385]" collectionCode="LACM" pageId="29" pageNumber="30" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
and
|
||
<materialsCitation box="[1180,1325,358,384]" collectionCode="RSM" pageId="29" pageNumber="30" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
, in contrast to the strongly curved form seen in juveniles such as
|
||
<materialsCitation box="[1088,1241,398,425]" collectionCode="CMNH" pageId="29" pageNumber="30" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
and BMRP 2002.4.1), are from 15 to 17 years old, and lack an EFS. The maximum number of changes in the entire growth series, 106, are seen in this growth category; 69 of the synontomorphies have a CI of 1.0 (
|
||
<tableCitation box="[621,709,518,544]" captionStart="Table 7" captionStartId="30.[525,580,248,269]" captionTargetBox="[516,1541,378,1912]" captionText="Table 7 Diagnostic characters of the subadult growth category of Tyrannosaurus rex. These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1)." pageId="29" pageNumber="30">Table 7</tableCitation>
|
||
). In the skull the changes are limited to the skull roof module. The maximum number of postcranial changes, 48, occur in this growth stage and affect the limb girdles and appendages. This category corresponds to growth stages 6 and 7 of the ontogram (
|
||
<figureCitation box="[631,697,637,663]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="29" pageNumber="30">Fig. 2</figureCitation>
|
||
) and to the first half of the exponential phase of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1327,1396,638,663]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1327,1396,638,663]" italics="true" pageId="29" pageNumber="30">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth curve (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[581,815,677,703]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[581,815,677,703]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[830,909,677,703]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="29" pageNumber="30">Fig. 12</figureCitation>
|
||
). The presence of medullary bone in
|
||
<materialsCitation box="[1355,1538,677,704]" collectionCode="BMRP" pageId="29" pageNumber="30" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[508,773,717,743]" journalOrPublisher="Science Advances" pageId="29" pageNumber="30" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[508,773,717,743]" italics="true" pageId="29" pageNumber="30">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
) marks the latest onset of sexual maturity (
|
||
<figureCitation box="[1290,1369,717,743]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="29" pageNumber="30">Fig. 12</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,233,1421]" pageId="29" pageNumber="30">
|
||
A mass estimate of 1,807 kg was published for
|
||
<materialsCitation box="[1063,1220,757,784]" collectionCode="LACM" pageId="29" pageNumber="30" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
by
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1260,1499,757,783]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1260,1499,757,783]" italics="true" pageId="29" pageNumber="30">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
; no other estimates have been published for this specimen. Therefore, a range of mass greater than ~1,300 kg and equal to or less than ~1,810 kg (the mass seen in young adults) is considered here as diagnostic of subadults.
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,233,1421]" pageId="29" pageNumber="30">
|
||
The transition from the long and low skull and narrow teeth of juveniles to the tall and powerful skull and thick teeth of subadults occurred shortly before the halfway point of life (assuming a lifespan of at least 28 years;
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1058,1292,996,1022]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1058,1292,996,1022]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
), within a narrow two-year interval between the growth categories, at the start of the exponential phase of growth (sensu
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[673,908,1075,1102]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[673,908,1075,1102]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
). This rapid change to the skull frame and dentition marks the onset of the increase in bite force; an order-of-magnitude increase in bite force between juvenile and adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[890,958,1156,1182]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[890,907,1156,1181]" italics="true" pageId="29" pageNumber="30">T</emphasis>
|
||
.
|
||
<emphasis box="[922,958,1156,1181]" italics="true" pageId="29" pageNumber="30">rex</emphasis>
|
||
</taxonomicName>
|
||
was found by
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[1134,1455,1155,1182]" journalOrPublisher="Biology Letters" pageId="29" pageNumber="30" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[1134,1455,1155,1182]" italics="true" pageId="29" pageNumber="30">Bates & Falkingham (2012)</emphasis>
|
||
</bibRefCitation>
|
||
. This increase is almost certainly congruent with the niche partitioning that is hypothesized to ecologically separate juveniles from adults (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1043,1459,1235,1261]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="29" pageNumber="30" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1043,1459,1235,1261]" italics="true" pageId="29" pageNumber="30">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
). Therefore, this extreme transition in skull shape and function occurs before the onset of somatic maturity (i.e., adult size; sensu
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[962,1196,1315,1341]" journalOrPublisher="Nature" pageId="29" pageNumber="30" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[962,1196,1315,1341]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) (
|
||
<figureCitation box="[1224,1303,1315,1341]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="29" pageNumber="30">Fig. 12</figureCitation>
|
||
). Also, the presence of medullary bone in
|
||
<materialsCitation box="[755,938,1354,1381]" collectionCode="BMRP" pageId="29" pageNumber="30" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
shows that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1081,1150,1355,1380]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="29" pageNumber="30" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1081,1150,1355,1380]" italics="true" pageId="29" pageNumber="30">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is typically reptilian given that sexual maturity precedes somatic maturity (
|
||
<bibRefCitation author="Erickson GM & Rogers KC & Varricchio DJ & Norell MA & Xu X." box="[1017,1251,1394,1420]" journalOrPublisher="Biology Letters" pageId="29" pageNumber="30" pagination="558 - 561" part="3" refId="ref53283" refString="Erickson GM, Rogers KC, Varricchio DJ, Norell MA, Xu X. 2007. Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition. Biology Letters 3 (5): 558 - 561 DOI 10.1098 / rsbl. 2007.0254." title="Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition" type="journal article" year="2007">
|
||
<emphasis box="[1017,1251,1394,1420]" italics="true" pageId="29" pageNumber="30">Erickson et al., 2007</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Lee AH & Werning S." box="[1266,1516,1394,1421]" journalOrPublisher="Proceedings of the National Association of Science" pageId="29" pageNumber="30" pagination="582 - 587" part="105" refId="ref54829" refString="Lee AH, Werning S. 2008. Sexual maturity in growing dinosaurs does not fit reptilian growth models. Proceedings of the National Association of Science 105 (2): 582 - 587 DOI 10.1073 / pnas. 0708903105." title="Sexual maturity in growing dinosaurs does not fit reptilian growth models" type="journal article" year="2008">
|
||
<emphasis box="[1266,1516,1394,1421]" italics="true" pageId="29" pageNumber="30">Lee & Werning, 2008</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,1459,1928]" box="[498,741,1459,1487]" pageId="29" pageNumber="30">
|
||
<heading bold="true" box="[498,741,1459,1487]" fontSize="11" level="3" pageId="29" pageNumber="30" reason="6">
|
||
<emphasis bold="true" box="[498,741,1459,1487]" italics="true" pageId="29" pageNumber="30">Skull architecture</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="29.[498,1542,1459,1928]" pageId="29" pageNumber="30">In the sixth growth stage, several changes to the lacrimal reflect an increase in bite force, including the 7-shaped bone, a rostrocaudally long ventral ramus, a dorsoventrally deep rostrodorsal process, a long rostroventral process that separates the nasal from the maxilla, a sliver-like maxillary process, the medially extended and convex medial margin of its dorsal ramus, the lengthened region caudal to the lacrimal pneumatic recess, and abruptly rostroventrally extending rostrodorsal margin of the rostral ramus.</paragraph>
|
||
<paragraph blockId="29.[498,1542,1459,1928]" lastBlockId="31.[498,1542,1513,1620]" lastPageId="31" lastPageNumber="32" pageId="29" pageNumber="30">Other architectural changes are seen in the lacrimal that are less obviously the result of high loads: the distal (=ventral) end of the orbitonasal ridge is mediolaterally wide but tapers ventrally; the ridge is not backswept; the orbitonasal ridge grades into the ventral ramus ahead of it; the caudal edge of the ridge is positioned far ahead of the caudal margin of the ventral ramus; the ridge extends abruptly rostroventrally dorsally, above the rostroventral ala; the rostrodorsal margin of the ventral ramus is straight or concave; and the rostroventral margin of the ventral ramus is straight. In the seventh growth stage, the rostroventral margin of ventral ramus of the lacrimal is concave.</paragraph>
|
||
<caption pageId="30" pageNumber="31" startId="30.[525,580,248,269]" targetBox="[516,1541,378,1912]" targetIsTable="true" targetPageId="30">
|
||
<paragraph blockId="30.[525,1515,248,358]" pageId="30" pageNumber="31">
|
||
<emphasis bold="true" box="[525,1392,248,270]" pageId="30" pageNumber="31">
|
||
Table 7 Diagnostic characters of the subadult growth category of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1199,1387,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1199,1387,249,270]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="30" pageNumber="31">
|
||
<table box="[516,1541,378,1912]" colsContinueFrom="31.[516,1524,290,1444]" gridcols="2" gridrows="41" pageId="30" pageNumber="31">
|
||
<tr box="[516,1541,378,400]" gridrow="0" pageId="30" pageNumber="31">
|
||
<th box="[516,1307,378,400]" gridcol="0" gridrow="0" pageId="30" pageNumber="31">Character</th>
|
||
<th box="[1372,1541,378,400]" gridcol="1" gridrow="0" pageId="30" pageNumber="31">
|
||
<emphasis bold="true" box="[1372,1524,378,400]" pageId="30" pageNumber="31">Growth change</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1541,419,442]" gridrow="1" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,419,442]" gridcol="0" gridrow="1" pageId="30" pageNumber="31">37. Lacrimal, bone, shape (PHYLO 65)</td>
|
||
<td box="[1372,1541,419,442]" gridcol="1" gridrow="1" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,457,479]" gridrow="2" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,457,479]" gridcol="0" gridrow="2" pageId="30" pageNumber="31">38. Lacrimal, cornual process, apex, shape (PHYLO 66)</td>
|
||
<td box="[1372,1541,457,479]" gridcol="1" gridrow="2" pageId="30" pageNumber="31">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1541,495,517]" gridrow="3" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,495,517]" gridcol="0" gridrow="3" pageId="30" pageNumber="31">40. Lacrimal, height above lacrimal pneumatic recess (PHYLO 69)</td>
|
||
<td box="[1372,1541,495,517]" gridcol="1" gridrow="3" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,532,554]" gridrow="4" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,532,554]" gridcol="0" gridrow="4" pageId="30" pageNumber="31">41. Lacrimal, rostral ramus, inflation (PHYLO 75)</td>
|
||
<td box="[1372,1541,532,554]" gridcol="1" gridrow="4" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,570,592]" gridrow="5" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,570,592]" gridcol="0" gridrow="5" pageId="30" pageNumber="31">41. Lacrimal, rostral ramus, inflation (PHYLO 75)</td>
|
||
<td box="[1372,1541,570,592]" gridcol="1" gridrow="5" pageId="30" pageNumber="31">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1541,607,630]" gridrow="6" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,607,630]" gridcol="0" gridrow="6" pageId="30" pageNumber="31">43. Lacrimal, number of recesses ahead of lacrimal pneumatic recess (PHYLO 78)</td>
|
||
<td box="[1372,1541,607,630]" gridcol="1" gridrow="6" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,645,667]" gridrow="7" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,645,667]" gridcol="0" gridrow="7" pageId="30" pageNumber="31">44. Lacrimal, medial pneumatic recess, presence (PHYLO 79)</td>
|
||
<td box="[1372,1541,645,667]" gridcol="1" gridrow="7" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,683,705]" gridrow="8" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,683,705]" gridcol="0" gridrow="8" pageId="30" pageNumber="31">45. Lacrimal, supraorbital ramus, length (PHYLO 82)</td>
|
||
<td box="[1372,1541,683,705]" gridcol="1" gridrow="8" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,720,742]" gridrow="9" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,720,742]" gridcol="0" gridrow="9" pageId="30" pageNumber="31">46. Lacrimal, supraorbital process, inflation (PHYLO 82)</td>
|
||
<td box="[1372,1541,720,742]" gridcol="1" gridrow="9" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,758,780]" gridrow="10" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,758,780]" gridcol="0" gridrow="10" pageId="30" pageNumber="31">219. Snout, dorsum, width caudal end</td>
|
||
<td box="[1372,1541,758,780]" gridcol="1" gridrow="10" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,795,817]" gridrow="11" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,795,817]" gridcol="0" gridrow="11" pageId="30" pageNumber="31">433. Lacrimal, dorsal ramus, medial edge, form</td>
|
||
<td box="[1372,1541,795,817]" gridcol="1" gridrow="11" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,833,855]" gridrow="12" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,833,855]" gridcol="0" gridrow="12" pageId="30" pageNumber="31">434. Lacrimal, medial fossa, rostral ridge, presence</td>
|
||
<td box="[1372,1541,833,855]" gridcol="1" gridrow="12" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,871,893]" gridrow="13" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,871,893]" gridcol="0" gridrow="13" pageId="30" pageNumber="31">436. Lacrimal, dorsal ramus, conchal surface, dorsal ridge, depth of caudal part</td>
|
||
<td box="[1372,1541,871,893]" gridcol="1" gridrow="13" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,908,930]" gridrow="14" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,908,930]" gridcol="0" gridrow="14" pageId="30" pageNumber="31">447. Lacrimal, dorsal ramus, region dorsal & rostrodorsal to lacrimal recess, form</td>
|
||
<td box="[1372,1541,908,930]" gridcol="1" gridrow="14" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,946,968]" gridrow="15" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,946,968]" gridcol="0" gridrow="15" pageId="30" pageNumber="31">454. Lacrimal, cornual process, lateral extent</td>
|
||
<td box="[1372,1541,946,968]" gridcol="1" gridrow="15" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,983,1005]" gridrow="16" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,983,1005]" gridcol="0" gridrow="16" pageId="30" pageNumber="31">455. Lacrimal, lacrimal pneumatic recess, foramen in ventral margin, presence</td>
|
||
<td box="[1372,1541,983,1005]" gridcol="1" gridrow="16" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1021,1043]" gridrow="17" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1021,1043]" gridcol="0" gridrow="17" pageId="30" pageNumber="31">461. Lacrimal, region caudal to lacrimal pneumatic recess, length & depth</td>
|
||
<td box="[1372,1541,1021,1043]" gridcol="1" gridrow="17" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1058,1081]" gridrow="18" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1058,1081]" gridcol="0" gridrow="18" pageId="30" pageNumber="31">462. Lacrimal, length lacrimal pneumatic recess relative to region behind it</td>
|
||
<td box="[1372,1541,1058,1081]" gridcol="1" gridrow="18" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1096,1118]" gridrow="19" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1096,1118]" gridcol="0" gridrow="19" pageId="30" pageNumber="31">465. Lacrimal, rostral ramus, subcutaneous surface height</td>
|
||
<td box="[1372,1541,1096,1118]" gridcol="1" gridrow="19" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1134,1156]" gridrow="20" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1134,1156]" gridcol="0" gridrow="20" pageId="30" pageNumber="31">466. Lacrimal, junction antorbital fossa & subcutaneous surface ahead of recess</td>
|
||
<td box="[1372,1541,1134,1156]" gridcol="1" gridrow="20" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1171,1193]" gridrow="21" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1171,1193]" gridcol="0" gridrow="21" pageId="30" pageNumber="31">468. Lacrimal, region between lacrimal pneumatic recess septum & distal recess</td>
|
||
<td box="[1372,1541,1171,1193]" gridcol="1" gridrow="21" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1209,1231]" gridrow="22" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1209,1231]" gridcol="0" gridrow="22" pageId="30" pageNumber="31">470. Lacrimal, pneumatic recess, proximity to maxilla</td>
|
||
<td box="[1372,1541,1209,1231]" gridcol="1" gridrow="22" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1247,1269]" gridrow="23" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1247,1269]" gridcol="0" gridrow="23" pageId="30" pageNumber="31">474. Lacrimal, rostral ramus, medial joint surfaces, height</td>
|
||
<td box="[1372,1541,1247,1269]" gridcol="1" gridrow="23" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1284,1306]" gridrow="24" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1284,1306]" gridcol="0" gridrow="24" pageId="30" pageNumber="31">476. Lacrimal, medial tab, dorsal margin</td>
|
||
<td box="[1372,1541,1284,1306]" gridcol="1" gridrow="24" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1322,1344]" gridrow="25" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1322,1344]" gridcol="0" gridrow="25" pageId="30" pageNumber="31">481. Lacrimal, maxillary process, height</td>
|
||
<td box="[1372,1541,1322,1344]" gridcol="1" gridrow="25" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1359,1381]" gridrow="26" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1359,1381]" gridcol="0" gridrow="26" pageId="30" pageNumber="31">482. Lacrimal, caudolateral shelf, form</td>
|
||
<td box="[1372,1541,1359,1381]" gridcol="1" gridrow="26" pageId="30" pageNumber="31">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1541,1397,1419]" gridrow="27" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1397,1419]" gridcol="0" gridrow="27" pageId="30" pageNumber="31">489. Lacrimal, frontal process, groove on dorsal surface, presence</td>
|
||
<td box="[1372,1541,1397,1419]" gridcol="1" gridrow="27" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1434,1457]" gridrow="28" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1434,1457]" gridcol="0" gridrow="28" pageId="30" pageNumber="31">517. Lacrimal, ventral ramus, proximal to distal ends</td>
|
||
<td box="[1372,1541,1434,1457]" gridcol="1" gridrow="28" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1472,1494]" gridrow="29" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1472,1494]" gridcol="0" gridrow="29" pageId="30" pageNumber="31">524. Lacrimal, ventral ramus, subcutaneous surface, caudal extension</td>
|
||
<td box="[1372,1541,1472,1494]" gridcol="1" gridrow="29" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1510,1532]" gridrow="30" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1510,1532]" gridcol="0" gridrow="30" pageId="30" pageNumber="31">1546. Scapulocoracoid, acromial region, lateral surface, form</td>
|
||
<td box="[1372,1541,1510,1532]" gridcol="1" gridrow="30" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1547,1569]" gridrow="31" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1547,1569]" gridcol="0" gridrow="31" pageId="30" pageNumber="31">1548. Scapula, glenoid, width joint surface relative to shaft</td>
|
||
<td box="[1372,1541,1547,1569]" gridcol="1" gridrow="31" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1585,1607]" gridrow="32" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1585,1607]" gridcol="0" gridrow="32" pageId="30" pageNumber="31">1549. Scapula, glenoid, anterolateral corner, form</td>
|
||
<td box="[1372,1541,1585,1607]" gridcol="1" gridrow="32" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1622,1644]" gridrow="33" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1622,1644]" gridcol="0" gridrow="33" pageId="30" pageNumber="31">1550. Scapula, lip of glenoid, emergence form ventral edge, form</td>
|
||
<td box="[1372,1541,1622,1644]" gridcol="1" gridrow="33" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1660,1682]" gridrow="34" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1660,1682]" gridcol="0" gridrow="34" pageId="30" pageNumber="31">1551. Scapula, acromion, medial surface, form</td>
|
||
<td box="[1372,1541,1660,1682]" gridcol="1" gridrow="34" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1698,1720]" gridrow="35" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1698,1720]" gridcol="0" gridrow="35" pageId="30" pageNumber="31">1561. Scapula, muscle scar along dorsal edge, distinctiveness</td>
|
||
<td box="[1372,1541,1698,1720]" gridcol="1" gridrow="35" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1735,1757]" gridrow="36" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1735,1757]" gridcol="0" gridrow="36" pageId="30" pageNumber="31">1568. Scapula, shaft, ventral surface adjacent to glenoid, orientation</td>
|
||
<td box="[1372,1541,1735,1757]" gridcol="1" gridrow="36" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1773,1795]" gridrow="37" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1773,1795]" gridcol="0" gridrow="37" pageId="30" pageNumber="31">1574. Coracoid, glenoid, orientation</td>
|
||
<td box="[1372,1541,1773,1795]" gridcol="1" gridrow="37" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1810,1832]" gridrow="38" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1810,1832]" gridcol="0" gridrow="38" pageId="30" pageNumber="31">1576. Coracoid, glenoid, separation from coracoid process, distinctiveness</td>
|
||
<td box="[1372,1541,1810,1832]" gridcol="1" gridrow="38" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1848,1870]" gridrow="39" pageId="30" pageNumber="31">
|
||
<td box="[516,1307,1848,1870]" gridcol="0" gridrow="39" pageId="30" pageNumber="31">1577. Coracoid, low ridge that extends anteriorly form biceps tubercle, presence</td>
|
||
<td box="[1372,1541,1848,1870]" gridcol="1" gridrow="39" pageId="30" pageNumber="31">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1541,1890,1912]" gridrow="40" pageId="30" pageNumber="31" rowspan-0="1">
|
||
<td box="[1372,1541,1890,1912]" gridcol="1" gridrow="40" pageId="30" pageNumber="31">(Continued)</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFDBFFB7FDEBFF7035F8FEB2" box="[525,738,248,270]" httpUri="http://table.plazi.org/id/DF5C84E2FFDBFFB7FDEBFF7035F8FEB2" pageId="31" pageNumber="32" startId="31.[525,580,248,269]" targetBox="[516,1524,290,1444]" targetIsTable="true" targetPageId="31">
|
||
<paragraph blockId="31.[525,738,248,270]" box="[525,738,248,270]" pageId="31" pageNumber="32">Table 7 (continued).</paragraph>
|
||
</caption>
|
||
<paragraph pageId="31" pageNumber="32">
|
||
<table box="[516,1524,290,1444]" colsContinueIn="30.[516,1541,378,1912]" gridcols="2" gridrows="31" pageId="31" pageNumber="32">
|
||
<tr box="[516,1524,290,312]" gridrow="0" pageId="31" pageNumber="32">
|
||
<th box="[516,1292,290,312]" gridcol="0" gridrow="0" pageId="31" pageNumber="32">Character</th>
|
||
<th box="[1372,1524,290,312]" gridcol="1" gridrow="0" pageId="31" pageNumber="32">
|
||
<emphasis bold="true" box="[1372,1524,290,312]" pageId="31" pageNumber="32">Growth change</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,332,354]" gridrow="1" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,332,354]" gridcol="0" gridrow="1" pageId="31" pageNumber="32">1581. Coracoid, ventral half of bone, form</td>
|
||
<td box="[1372,1524,332,354]" gridcol="1" gridrow="1" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,369,391]" gridrow="2" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,369,391]" gridcol="0" gridrow="2" pageId="31" pageNumber="32">1583. Coracoid process, length, depth & curvature</td>
|
||
<td box="[1372,1524,369,391]" gridcol="1" gridrow="2" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,407,429]" gridrow="3" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,407,429]" gridcol="0" gridrow="3" pageId="31" pageNumber="32">1584. Coracoid, medial fossa between coracoid foramen & anterior edge of bone</td>
|
||
<td box="[1372,1524,407,429]" gridcol="1" gridrow="3" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,445,467]" gridrow="4" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,445,467]" gridcol="0" gridrow="4" pageId="31" pageNumber="32">1707. D II, PH 2, proximal joint surface, ventral margin, form</td>
|
||
<td box="[1372,1524,445,467]" gridcol="1" gridrow="4" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,482,504]" gridrow="5" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,482,504]" gridcol="0" gridrow="5" pageId="31" pageNumber="32">1711. D II, PH 2, lateral distal condyle, orientation, lateral view</td>
|
||
<td box="[1372,1524,482,504]" gridcol="1" gridrow="5" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,520,542]" gridrow="6" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,520,542]" gridcol="0" gridrow="6" pageId="31" pageNumber="32">1712. D II, PH 2, medial distal condyle, elevation</td>
|
||
<td box="[1372,1524,520,542]" gridcol="1" gridrow="6" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,557,579]" gridrow="7" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,557,579]" gridcol="0" gridrow="7" pageId="31" pageNumber="32">1714. D II, PH 2, proximodorsal flange, width</td>
|
||
<td box="[1372,1524,557,579]" gridcol="1" gridrow="7" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,595,617]" gridrow="8" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,595,617]" gridcol="0" gridrow="8" pageId="31" pageNumber="32">1715. D II, PH 2, medial distal condyle, medial margin, orientation</td>
|
||
<td box="[1372,1524,595,617]" gridcol="1" gridrow="8" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,632,654]" gridrow="9" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,632,654]" gridcol="0" gridrow="9" pageId="31" pageNumber="32">1719. D II, PH 2, medial distal condyle, ventral extent</td>
|
||
<td box="[1372,1524,632,654]" gridcol="1" gridrow="9" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,670,692]" gridrow="10" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,670,692]" gridcol="0" gridrow="10" pageId="31" pageNumber="32">1748. D III, PH 1, proximal joint surface, form</td>
|
||
<td box="[1372,1524,670,692]" gridcol="1" gridrow="10" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,708,730]" gridrow="11" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,708,730]" gridcol="0" gridrow="11" pageId="31" pageNumber="32">1754. D III, PH 1, lateral caudolateral ligament pit, frm</td>
|
||
<td box="[1372,1524,708,730]" gridcol="1" gridrow="11" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,745,767]" gridrow="12" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,745,767]" gridcol="0" gridrow="12" pageId="31" pageNumber="32">1779. D IV, PH 1, supracondylar pit, form</td>
|
||
<td box="[1372,1524,745,767]" gridcol="1" gridrow="12" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,783,805]" gridrow="13" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,783,805]" gridcol="0" gridrow="13" pageId="31" pageNumber="32">1782. D IV, PH 1, ventromedial condyle, angle relative to shaft</td>
|
||
<td box="[1372,1524,783,805]" gridcol="1" gridrow="13" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,821,843]" gridrow="14" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,821,843]" gridcol="0" gridrow="14" pageId="31" pageNumber="32">1784. D IV, PH 1, proximal joint surface, ventral notch, mediolateral position</td>
|
||
<td box="[1372,1524,821,843]" gridcol="1" gridrow="14" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,858,880]" gridrow="15" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,858,880]" gridcol="0" gridrow="15" pageId="31" pageNumber="32">1787. D IV, PH 1, distal joint surface, proportions</td>
|
||
<td box="[1372,1524,858,880]" gridcol="1" gridrow="15" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,896,918]" gridrow="16" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,896,918]" gridcol="0" gridrow="16" pageId="31" pageNumber="32">1790. D IV, PH 1, scar on shaft ahead of proximal end, prominence</td>
|
||
<td box="[1372,1524,896,918]" gridcol="1" gridrow="16" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,933,955]" gridrow="17" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,933,955]" gridcol="0" gridrow="17" pageId="31" pageNumber="32">1792. D IV, PH 1, dorsolateral surface, form</td>
|
||
<td box="[1372,1524,933,955]" gridcol="1" gridrow="17" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,971,993]" gridrow="18" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,971,993]" gridcol="0" gridrow="18" pageId="31" pageNumber="32">1793. D IV, PH 1, shaft, ridge lateral to supracondylar pit, form</td>
|
||
<td box="[1372,1524,971,993]" gridcol="1" gridrow="18" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1008,1030]" gridrow="19" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1008,1030]" gridcol="0" gridrow="19" pageId="31" pageNumber="32">1806. D IV, PH 3, form</td>
|
||
<td box="[1372,1524,1008,1030]" gridcol="1" gridrow="19" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1046,1068]" gridrow="20" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1046,1068]" gridcol="0" gridrow="20" pageId="31" pageNumber="32">1809. D IV, PH 3, ratio of mediolateral width to dorsoventral height</td>
|
||
<td box="[1372,1524,1046,1068]" gridcol="1" gridrow="20" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1084,1106]" gridrow="21" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1084,1106]" gridcol="0" gridrow="21" pageId="31" pageNumber="32">1810. D IV, PH 3, proximal joint surface, medial margin, form</td>
|
||
<td box="[1372,1524,1084,1106]" gridcol="1" gridrow="21" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1121,1143]" gridrow="22" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1121,1143]" gridcol="0" gridrow="22" pageId="31" pageNumber="32">1812. D IV, PH 3, medial collateral ligament point, depth</td>
|
||
<td box="[1372,1524,1121,1143]" gridcol="1" gridrow="22" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1159,1181]" gridrow="23" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1159,1181]" gridcol="0" gridrow="23" pageId="31" pageNumber="32">1814. D IV, PH 3, distal condyles, divergence</td>
|
||
<td box="[1372,1524,1159,1181]" gridcol="1" gridrow="23" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1196,1218]" gridrow="24" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1196,1218]" gridcol="0" gridrow="24" pageId="31" pageNumber="32">1815. D IV, PH 3, flexor muscle scar, form</td>
|
||
<td box="[1372,1524,1196,1218]" gridcol="1" gridrow="24" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1234,1256]" gridrow="25" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1234,1256]" gridcol="0" gridrow="25" pageId="31" pageNumber="32">1817. D IV PH 4, dorsum, form</td>
|
||
<td box="[1372,1524,1234,1256]" gridcol="1" gridrow="25" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1272,1294]" gridrow="26" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1272,1294]" gridcol="0" gridrow="26" pageId="31" pageNumber="32">1818. D IV PH 4, dorsum, mediolateral orientation</td>
|
||
<td box="[1372,1524,1272,1294]" gridcol="1" gridrow="26" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1309,1331]" gridrow="27" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1309,1331]" gridcol="0" gridrow="27" pageId="31" pageNumber="32">1820. D IV PH 4, distal joint surface, form</td>
|
||
<td box="[1372,1524,1309,1331]" gridcol="1" gridrow="27" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1347,1369]" gridrow="28" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1347,1369]" gridcol="0" gridrow="28" pageId="31" pageNumber="32">1824. D IV PH 4, distal joint surface, dorsal margin, indentation, presence</td>
|
||
<td box="[1372,1524,1347,1369]" gridcol="1" gridrow="28" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1384,1406]" gridrow="29" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1384,1406]" gridcol="0" gridrow="29" pageId="31" pageNumber="32">1826. D IV PH 4, distal condylar region, posterior margin, undercut</td>
|
||
<td box="[1372,1524,1384,1406]" gridcol="1" gridrow="29" pageId="31" pageNumber="32">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1422,1444]" gridrow="30" pageId="31" pageNumber="32">
|
||
<td box="[516,1292,1422,1444]" gridcol="0" gridrow="30" pageId="31" pageNumber="32">1848. Growth rings, number</td>
|
||
<td box="[1372,1524,1422,1444]" gridcol="1" gridrow="30" pageId="31" pageNumber="32">2 ==> 3</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="31.[498,1542,1658,1888]" box="[498,695,1658,1686]" pageId="31" pageNumber="32">
|
||
<heading bold="true" box="[498,695,1658,1686]" fontSize="11" level="3" pageId="31" pageNumber="32" reason="6">
|
||
<emphasis bold="true" box="[498,695,1658,1686]" italics="true" pageId="31" pageNumber="32">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="31.[498,1542,1658,1888]" lastBlockId="32.[498,1541,234,301]" lastPageId="32" lastPageNumber="33" pageId="31" pageNumber="32">In the sixth growth stage, several changes to the lacrimal are seen, including the joint surface for the prefrontal on the dorsal ramus of the lacrimal is a dorsoventrally deep and laterally shallow groove that caudally twists to face mediodorsally, the joint surface for the nasal on the rostral ramus covers its ventral third, the dorsal half of the medial joint surfaces is dorsoventrally shallow, the medial joint surface (behind the medial tube) is concave in vertical section, a groove on the dorsal surface of the frontal process is present, and a medially extending process behind the joint surface for the prefrontal is present.</paragraph>
|
||
<paragraph blockId="32.[498,1541,339,489]" box="[498,809,339,367]" pageId="32" pageNumber="33">
|
||
<heading bold="true" box="[498,809,339,367]" fontSize="11" level="3" pageId="32" pageNumber="33" reason="6">
|
||
<emphasis bold="true" box="[498,809,339,367]" italics="true" pageId="32" pageNumber="33">Subcutaneous surface</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="32.[498,1541,339,489]" pageId="32" pageNumber="33">In the sixth growth stage, the subcutaneous surface of the lacrimal wraps onto the caudal surface of the ventral ramus and the coarse patch caudodorsal to the lacrimal pneumatic recess is absent.</paragraph>
|
||
<paragraph blockId="32.[498,1542,528,598]" box="[498,828,528,556]" pageId="32" pageNumber="33">
|
||
<heading bold="true" box="[498,828,528,556]" fontSize="11" level="3" pageId="32" pageNumber="33" reason="6">
|
||
<emphasis bold="true" box="[498,828,528,556]" italics="true" pageId="32" pageNumber="33">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="32.[498,1542,528,598]" box="[498,1542,572,598]" pageId="32" pageNumber="33">In the sixth growth stage, the low ridge caudodorsal to the lacrimal pneumatic recess is lost.</paragraph>
|
||
<paragraph blockId="32.[498,1542,637,1703]" box="[498,822,637,665]" pageId="32" pageNumber="33">
|
||
<heading bold="true" box="[498,822,637,665]" fontSize="11" level="3" pageId="32" pageNumber="33" reason="6">
|
||
<emphasis bold="true" box="[498,822,637,665]" italics="true" pageId="32" pageNumber="33">Paranasal pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="32.[498,1542,637,1703]" pageId="32" pageNumber="33">In the sixth growth stage, the invasive antorbital air sac produces the medial pneumatic recess of the lacrimal. Some pneumatic changes are almost certainly an epiphenomenon of the overall dorsoventral and mediolateral expansion of the skull, including the distal pneumatic recess of the lacrimal that is positioned close to maxilla. Some pneumatic changes more directly reflect the resorptive tendency of pneumatic epithelia, including the absorption of the caudoventral margin of the medial pneumatic recess.</paragraph>
|
||
<paragraph blockId="32.[498,1542,637,1703]" pageId="32" pageNumber="33">Inflation of the dorsal ramus of the lacrimal has several simultaneous effects upon the bone, including obscuring the apex of the cornual process and the process in general; increase in the height of the cornual process; loss of the ridge that rostrally bounds the conchal surface; increase of the depth of the ridge above the conchal surface; the convex caudodorsal region of the shallow conchal surface; the convex lateral surface around the lacrimal pneumatic recess; the laterally bulging region of the cornual process of the lacrimal; increase in the height of the subcutaneous surface above the antorbital fossa to twice the height of the rostral ramus of the lacrimal; reduction of the size of the lacrimal pneumatic recess to less than half the length of the region behind it; inflation of the region below the lacrimal pneumatic recess, and the septum ahead of it, which merges the junction of the antorbital fossa and subcutaneous surface ahead of the lacrimal pneumatic recess; the shallowly concave region between the distal recess and the septum ahead of the lacrimal pneumatic recess; the lengthened septum between the accessory recesses; the shallowly concave distal pneumatic recess of the lacrimal; the widened supraorbital process and reduction of its length and increase in its height; and the caudal end of the snout is widened by the inflation of the lacrimals.</paragraph>
|
||
<paragraph blockId="32.[498,1542,637,1703]" pageId="32" pageNumber="33">In the seventh growth stage, inflation has obliterated the cornual process of the lacrimal. A single accessory recess, the distal recess, is present ahead of the lacrimal pneumatic recess, which is reduced to a foramen.</paragraph>
|
||
<paragraph blockId="32.[498,1525,1742,1891]" box="[498,741,1742,1770]" pageId="32" pageNumber="33">
|
||
<heading bold="true" box="[498,741,1742,1770]" fontSize="11" level="3" pageId="32" pageNumber="33" reason="6">
|
||
<emphasis bold="true" box="[498,741,1742,1770]" italics="true" pageId="32" pageNumber="33">Neurovasculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="32.[498,1525,1742,1891]" pageId="32" pageNumber="33">In the sixth growth stage, the foramen in the ridge that bounds the lacrimal pneumatic recess is absent, and the dorsal margin of the medial tube of the lacrimal extends rostrodorsally.</paragraph>
|
||
<paragraph blockId="33.[498,1542,233,1777]" box="[498,808,233,261]" pageId="33" pageNumber="34">
|
||
<heading bold="true" box="[498,808,233,261]" fontSize="11" level="3" pageId="33" pageNumber="34" reason="6">
|
||
<emphasis bold="true" box="[498,808,233,261]" italics="true" pageId="33" pageNumber="34">Appendicular skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="33.[498,1542,233,1777]" pageId="33" pageNumber="34">In the seventh growth stage several changes are seen in the pectoral girdle and limb. In the scapula, the acromial region is most deeply concave on the scapula, the glenoid fossa has an intermediate position between lateral and ventral, the glenoid fossa is narrower than the shaft, the anterolateral corner of glenoid widens toward the coracoid, the medial surface of the acromion is convex, the muscle scar along the dorsal edge of the shaft is indistinct, and the ridge on lateral surface of the shaft is absent. The coracoid also shows changes, including a posteroventral orientation of the glenoid, an abrupt separation of the glenoid from the coracoid process, loss of the ridge that extends anteriorly from the biceps tubercle, the ventral half of the bone is convex, and the medial fossa ahead of the coracoid foramen stops short of the anterior edge of the bone. Finally, the deltoid scar at the distal end of the deltopectoral crest of the humerus is deeply inset.</paragraph>
|
||
<paragraph blockId="33.[498,1542,233,1777]" pageId="33" pageNumber="34">In the seventh growth stage, the lip of the glenoid of the scapula extends from the shaft at an abrupt angle, the shaft is not distinctly narrow between the acromial region and the shaft, and the ventral surface of the shaft next to the glenoid faces ventrally. Changes in the coracoid include the anterolateral orientation of the coracoid foramen and the stout form of the coracoid process.</paragraph>
|
||
<paragraph blockId="33.[498,1542,233,1777]" pageId="33" pageNumber="34">The changes to the pelvic limb happen in the pes. In the sixth growth stage, the ventral margin of the proximal joint surface of D II PH 2 is concave, the anterior end of the lateral distal condyle is flattened, the medial distal condyle extends anterodorsally, the proximodorsal flange is wide, the medial distal condyle extends anterolaterally, and it extends posteriorly below the level of the posterior margin of the medial collateral ligament pit. In D III PH 1, the proximal joint surface is dissected by sulci and the lateral collateral ligament pit is lens-shaped. In D IV PH 1, the ventromedial condyle extends at a steep angle relative to the shaft, the ventral notch of the proximal joint surface is positioned at the midline, the distal joint surface is much wider than tall, the scar on the lateral surface of the shaft ahead of the proximal end is distinct, the dorsolateral surface of the shaft is convex, and the ridge lateral to the supracondylar pit is short. In D IV PH 3, the phalanx is short, the proximal joint surface is distinctly wider than tall, the lateral collateral ligament pit is deep, and the flexor muscle scar is coarse. In D IV PH 4, the dorsum of the bone is wide, the mediolateral orientation of the dorsum is distinct, the proximal end of the bone is wider than the distal end, the distal joint surface is distinctly concave, the entire posterior margin of the distal condylar region is undercut by a groove that separates it from the ventral surface of the shaft, and the indentation along the dorsal margin of the distal joint surface is wide.</paragraph>
|
||
<paragraph blockId="33.[498,1542,233,1777]" pageId="33" pageNumber="34">In the seventh growth stage, changes are seen in D IV PH 1, where the supracondylar pit is a deep crease that sharply elevates the distal condyles. In D IV PH 3, supracondylar pit is deep, the medial margin of the proximal joint surface is distinctly convex, the medial collateral ligament pit is large, and the distal condyles shallowly diverge from each other.</paragraph>
|
||
<paragraph blockId="33.[498,1542,1822,1934]" box="[498,697,1822,1852]" pageId="33" pageNumber="34">
|
||
<heading bold="true" box="[498,697,1822,1852]" fontSize="12" level="2" pageId="33" pageNumber="34" reason="0">
|
||
<emphasis bold="true" box="[498,697,1822,1852]" pageId="33" pageNumber="34">Young adults</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="33.[498,1542,1822,1934]" lastBlockId="34.[498,1542,1569,1915]" lastPageId="34" lastPageNumber="35" pageId="33" pageNumber="34">
|
||
Young adults (=subadult of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[826,1057,1868,1894]" journalOrPublisher="Nature" pageId="33" pageNumber="34" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[826,1057,1868,1894]" italics="true" pageId="33" pageNumber="34">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) have a skull length from ~1.16 to 1.4 m, tall skulls with a length to height ratio of 2.6
|
||
<emphasis box="[1011,1026,1909,1934]" italics="true" pageId="33" pageNumber="34">–</emphasis>
|
||
2.3, are from 18 to 22 years old, lack an EFS, and, as seen in subadults, they initially have a high number of synontomorphies, which thereafter drops in frequency (
|
||
<figureCitation box="[860,936,1609,1635]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="34" pageNumber="35">Figs. 2</figureCitation>
|
||
and 4). In total, this growth stage has 77 growth changes; 29 have a CI of 1.0 (
|
||
<tableCitation box="[840,926,1649,1675]" captionStart="Table 8" captionStartId="34.[525,580,248,269]" captionTargetBox="[516,1524,378,1494]" captionText="Table 8 Diagnostic characters of the young adult growth category of Tyrannosaurus rex. These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1)." httpUri="http://table.plazi.org/id/DF5C84E2FFE6FF8AFDEBFF70329BFEDA" pageId="34" pageNumber="35" tableUuid="DF5C84E2FFE6FF8AFDEBFF70329BFEDA">Table 8</tableCitation>
|
||
). In the skull, the greatest number of changes is seen in the skull roof and braincase modules. In contrast to subadults, growth changes are seen across every module of the skull. Postcranial changes are limited to the hindlimb. This category corresponds to growth stages 8
|
||
<emphasis box="[1018,1033,1769,1794]" italics="true" pageId="34" pageNumber="35">–</emphasis>
|
||
12 of the ontogram (
|
||
<figureCitation box="[1271,1334,1769,1795]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="34" pageNumber="35">Fig. 2</figureCitation>
|
||
), occurring in the last half of the exponential phase of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[960,1027,1809,1834]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[960,1027,1809,1834]" italics="true" pageId="34" pageNumber="35">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth curve (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1205,1434,1808,1835]" journalOrPublisher="Nature" pageId="34" pageNumber="35" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1205,1434,1808,1835]" italics="true" pageId="34" pageNumber="35">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1448,1525,1808,1835]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="34" pageNumber="35">Fig. 12</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFE6FF8AFDEBFF70329BFEDA" httpUri="http://table.plazi.org/id/DF5C84E2FFE6FF8AFDEBFF70329BFEDA" pageId="34" pageNumber="35" startId="34.[525,580,248,269]" targetBox="[516,1524,378,1494]" targetIsTable="true" targetPageId="34">
|
||
<paragraph blockId="34.[525,1515,248,358]" pageId="34" pageNumber="35">
|
||
<emphasis bold="true" box="[525,1449,248,270]" pageId="34" pageNumber="35">
|
||
Table 8 Diagnostic characters of the young adult growth category of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1253,1443,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1253,1443,249,270]" italics="true" pageId="34" pageNumber="35">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. PHYLO, corresponding homologous phylogenetic character (see Data S1).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="34" pageNumber="35">
|
||
<table box="[516,1524,378,1494]" gridcols="2" gridrows="30" pageId="34" pageNumber="35">
|
||
<tr box="[516,1524,378,400]" gridrow="0" pageId="34" pageNumber="35">
|
||
<th box="[516,1235,378,400]" gridcol="0" gridrow="0" pageId="34" pageNumber="35">Character</th>
|
||
<th box="[1372,1524,378,400]" gridcol="1" gridrow="0" pageId="34" pageNumber="35">
|
||
<emphasis bold="true" box="[1372,1524,378,400]" pageId="34" pageNumber="35">Growth change</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,419,442]" gridrow="1" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,419,442]" gridcol="0" gridrow="1" pageId="34" pageNumber="35">18. Maxilla, interfenestral strut, length (PHYLO 31)</td>
|
||
<td box="[1372,1524,419,442]" gridcol="1" gridrow="1" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,457,479]" gridrow="2" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,457,479]" gridcol="0" gridrow="2" pageId="34" pageNumber="35">24. Maxilla, alveolar process, subcutaneous surface, fossae (PHYLO 44)</td>
|
||
<td box="[1372,1524,457,479]" gridcol="1" gridrow="2" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,495,517]" gridrow="3" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,495,517]" gridcol="0" gridrow="3" pageId="34" pageNumber="35">109. Otoccipital, paroccipital process, caudal surface, form (PHYLO 200)</td>
|
||
<td box="[1372,1524,495,517]" gridcol="1" gridrow="3" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,532,554]" gridrow="4" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,532,554]" gridcol="0" gridrow="4" pageId="34" pageNumber="35">253. Premaxilla, lateral margin, form</td>
|
||
<td box="[1372,1524,532,554]" gridcol="1" gridrow="4" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,570,592]" gridrow="5" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,570,592]" gridcol="0" gridrow="5" pageId="34" pageNumber="35">376. Maxilla, antorbital fossa, mediolateral depth</td>
|
||
<td box="[1372,1524,570,592]" gridcol="1" gridrow="5" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,607,629]" gridrow="6" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,607,629]" gridcol="0" gridrow="6" pageId="34" pageNumber="35">523. Lacrimal, subcutaneous surface, texture</td>
|
||
<td box="[1372,1524,607,629]" gridcol="1" gridrow="6" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,645,667]" gridrow="7" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,645,667]" gridcol="0" gridrow="7" pageId="34" pageNumber="35">651. Postorbital, jugal ramus, width, caudal</td>
|
||
<td box="[1372,1524,645,667]" gridcol="1" gridrow="7" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,683,705]" gridrow="8" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,683,705]" gridcol="0" gridrow="8" pageId="34" pageNumber="35">717. Squamosal, dorsotemporal fossa, lateral half, texture</td>
|
||
<td box="[1372,1524,683,705]" gridcol="1" gridrow="8" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,720,742]" gridrow="9" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,720,742]" gridcol="0" gridrow="9" pageId="34" pageNumber="35">737. Quadratojugal, squamosal process, ridge, width</td>
|
||
<td box="[1372,1524,720,742]" gridcol="1" gridrow="9" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,758,780]" gridrow="10" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,758,780]" gridcol="0" gridrow="10" pageId="34" pageNumber="35">741. Quadratojugal, dorsal joint surface for quadratojugal, texture & depth</td>
|
||
<td box="[1372,1524,758,780]" gridcol="1" gridrow="10" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,795,817]" gridrow="11" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,795,817]" gridcol="0" gridrow="11" pageId="34" pageNumber="35">742. Quadratojugal, dorsal joint surface for quadrate, rostral margin, form</td>
|
||
<td box="[1372,1524,795,817]" gridcol="1" gridrow="11" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,833,855]" gridrow="12" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,833,855]" gridcol="0" gridrow="12" pageId="34" pageNumber="35">882. Frontals, proportions apposed bones</td>
|
||
<td box="[1372,1524,833,855]" gridcol="1" gridrow="12" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,871,893]" gridrow="13" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,871,893]" gridcol="0" gridrow="13" pageId="34" pageNumber="35">913. Frontal, joint surface for the prefrontal, rostrocaudal position</td>
|
||
<td box="[1372,1524,871,893]" gridcol="1" gridrow="13" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,908,930]" gridrow="14" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,908,930]" gridcol="0" gridrow="14" pageId="34" pageNumber="35">931. Frontal, orbital surface, medial margin, orientation</td>
|
||
<td box="[1372,1524,908,930]" gridcol="1" gridrow="14" pageId="34" pageNumber="35">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,946,968]" gridrow="15" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,946,968]" gridcol="0" gridrow="15" pageId="34" pageNumber="35">1014. Otoccipital, metotic strut, oval scar complex, participation</td>
|
||
<td box="[1372,1524,946,968]" gridcol="1" gridrow="15" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,983,1005]" gridrow="16" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,983,1005]" gridcol="0" gridrow="16" pageId="34" pageNumber="35">1017. Subcondylar recesses, distance from each other</td>
|
||
<td box="[1372,1524,983,1005]" gridcol="1" gridrow="16" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1021,1043]" gridrow="17" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1021,1043]" gridcol="0" gridrow="17" pageId="34" pageNumber="35">1043. Basisphenoid, oval scar, texture</td>
|
||
<td box="[1372,1524,1021,1043]" gridcol="1" gridrow="17" pageId="34" pageNumber="35">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,1059,1081]" gridrow="18" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1059,1081]" gridcol="0" gridrow="18" pageId="34" pageNumber="35">1048. Basisphenoid, spout, presence</td>
|
||
<td box="[1372,1524,1059,1081]" gridcol="1" gridrow="18" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1096,1118]" gridrow="19" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1096,1118]" gridcol="0" gridrow="19" pageId="34" pageNumber="35">1052. Basisphenoid, flange form ventral end of preotic pendant</td>
|
||
<td box="[1372,1524,1096,1118]" gridcol="1" gridrow="19" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1134,1156]" gridrow="20" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1134,1156]" gridcol="0" gridrow="20" pageId="34" pageNumber="35">1055. Supraoccipital, bar across lateral processes, prominence</td>
|
||
<td box="[1372,1524,1134,1156]" gridcol="1" gridrow="20" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1171,1193]" gridrow="21" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1171,1193]" gridcol="0" gridrow="21" pageId="34" pageNumber="35">1082. Dentary, chin, subcutaneous surface, texture</td>
|
||
<td box="[1372,1524,1171,1193]" gridcol="1" gridrow="21" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1209,1231]" gridrow="22" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1209,1231]" gridcol="0" gridrow="22" pageId="34" pageNumber="35">1111. Dentary, ventral bar, depth below Meckelian fossa</td>
|
||
<td box="[1372,1524,1209,1231]" gridcol="1" gridrow="22" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1247,1269]" gridrow="23" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1247,1269]" gridcol="0" gridrow="23" pageId="34" pageNumber="35">1168. Angular, 2ndry ridge, form</td>
|
||
<td box="[1372,1524,1247,1269]" gridcol="1" gridrow="23" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1284,1306]" gridrow="24" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1284,1306]" gridcol="0" gridrow="24" pageId="34" pageNumber="35">1647. Fibula, ratio of midheight length to total height</td>
|
||
<td box="[1372,1524,1284,1306]" gridcol="1" gridrow="24" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1322,1344]" gridrow="25" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1322,1344]" gridcol="0" gridrow="25" pageId="34" pageNumber="35">1657. Fibula, proximal end, dorsal margin, form</td>
|
||
<td box="[1372,1524,1322,1344]" gridcol="1" gridrow="25" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1359,1381]" gridrow="26" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1359,1381]" gridcol="0" gridrow="26" pageId="34" pageNumber="35">1672. Fibula, anterior surface proximal surface to bipartite scar</td>
|
||
<td box="[1372,1524,1359,1381]" gridcol="1" gridrow="26" pageId="34" pageNumber="35">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,1397,1419]" gridrow="27" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1397,1419]" gridcol="0" gridrow="27" pageId="34" pageNumber="35">1675. Fibula, medial fossa, differentiation</td>
|
||
<td box="[1372,1524,1397,1419]" gridcol="1" gridrow="27" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1434,1457]" gridrow="28" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1434,1457]" gridcol="0" gridrow="28" pageId="34" pageNumber="35">16.83. Fibula, ventral end below bipartite scar, form</td>
|
||
<td box="[1372,1524,1434,1457]" gridcol="1" gridrow="28" pageId="34" pageNumber="35">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,1472,1494]" gridrow="29" pageId="34" pageNumber="35">
|
||
<td box="[516,1235,1472,1494]" gridcol="0" gridrow="29" pageId="34" pageNumber="35">1848. Growth rings, number</td>
|
||
<td box="[1372,1524,1472,1494]" gridcol="1" gridrow="29" pageId="34" pageNumber="35">3 ==> 4</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="34.[498,1542,1569,1915]" lastBlockId="35.[498,1542,234,540]" lastPageId="35" lastPageNumber="36" pageId="34" pageNumber="35">
|
||
Mass estimates for the young adult
|
||
<materialsCitation box="[947,1182,1848,1875]" collectionCode="TMP" pageId="34" pageNumber="35" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
range from 3,230 kg (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" journalOrPublisher="Nature" pageId="34" pageNumber="35" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="34" pageNumber="35">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) to 4,469 kg (
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[794,1179,1888,1915]" journalOrPublisher="Anatomical Record" pageId="34" pageNumber="35" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[794,1179,1888,1915]" italics="true" pageId="34" pageNumber="35">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
). Another young adult,
|
||
<materialsCitation box="[498,631,234,261]" collectionCode="MOR" pageId="35" pageNumber="36" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, has been estimated at 6,100 kg (
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[1023,1409,234,261]" journalOrPublisher="Anatomical Record" pageId="35" pageNumber="36" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[1023,1409,234,261]" italics="true" pageId="35" pageNumber="36">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
). This doubling in mass from 1,810 kg subadults to 3,000 kg young adults occurred halfway through life, in the 3-year interval (15
|
||
<emphasis box="[937,952,315,340]" italics="true" pageId="35" pageNumber="36">–</emphasis>
|
||
18 years) in the middle of the exponential phase of growth (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[600,834,354,380]" journalOrPublisher="Nature" pageId="35" pageNumber="36" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[600,834,354,380]" italics="true" pageId="35" pageNumber="36">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
); this ontogenetic increase in mass corresponds to the phylogenetic threshold in mass, where
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[958,1027,394,419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[958,1027,394,419]" italics="true" pageId="35" pageNumber="36">T. rex</emphasis>
|
||
</taxonomicName>
|
||
exceeds the 3,000 kg limit of its closest relatives, namely
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[703,968,433,459]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[703,968,433,459]" italics="true" pageId="35" pageNumber="36">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[1030,1138,433,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1030,1138,433,459]" italics="true" pageId="35" pageNumber="36">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[1157,1375,433,460]" journalOrPublisher="PeerJ" pageId="35" pageNumber="36" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[1157,1375,433,460]" italics="true" pageId="35" pageNumber="36">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
). Therefore, this character state in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[760,829,474,499]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[760,829,474,499]" italics="true" pageId="35" pageNumber="36">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is hypermorphic in that it suprasses the ancestral condition (cf.
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[544,779,513,539]" journalOrPublisher="Nature" pageId="35" pageNumber="36" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[544,779,513,539]" italics="true" pageId="35" pageNumber="36">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="35.[498,1542,578,1406]" box="[498,741,578,606]" pageId="35" pageNumber="36">
|
||
<heading bold="true" box="[498,741,578,606]" fontSize="11" level="3" pageId="35" pageNumber="36" reason="6">
|
||
<emphasis bold="true" box="[498,741,578,606]" italics="true" pageId="35" pageNumber="36">Skull architecture</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="35.[498,1542,578,1406]" pageId="35" pageNumber="36">In the seventh growth stage, the rostral margin of the rostroventral lamina of the lacrimal is convex, the jugal ramus of the postorbital is rostrocaudally wide and parallel-sided, the apposed frontals are wider than long, the caudal end of the joint surface for the prefrontal on the frontal is positioned below the joint surface for the postorbital, the orbital surface of the frontal is horizontally oriented, the midline strut of the nuchal crest that is above the supraoccipital is prominent, the ventral bar of the dentary that is below the Meckelian groove is dorsoventrally deep, and the secondary ridge on the medial surface of the angular is distinct.</paragraph>
|
||
<paragraph blockId="35.[498,1542,578,1406]" pageId="35" pageNumber="36">In the eighth growth stage, the dorsal and ventral margins of the ascending ramus of the maxilla converge as they extend caudally, the point of rostralmost incursion of the internal antorbital fenestra is positioned within less than five teeth of the caudal end of the tooth row, the subocular process of the postorbital is as long or longer than the frontal process, and the chin of the dentary is positioned ahead of the fourth alveolus.</paragraph>
|
||
<paragraph blockId="35.[498,1542,578,1406]" pageId="35" pageNumber="36">In the ninth growth stage, the caudal end of the antorbital fossa has a uniform height below the internal antorbital fenestra and it is obliterated (i.e., fades out) ahead of the jugal ramus, and the lateral margin of the premaxilla is straight or convex. In the tenth growth stage, the concavity in the caudal surface of the nuchal crest fades below the dorsal margin of the crest, and the ventral surface of the neck of the occipital condyle has the form of a deep pit between pillars. In the eleventh growth stage, the dorsal surface of the neck of the occipital condyle is penetrated by a shallow pit.</paragraph>
|
||
<paragraph blockId="35.[498,1542,1444,1913]" box="[498,695,1444,1472]" pageId="35" pageNumber="36">
|
||
<heading bold="true" box="[498,695,1444,1472]" fontSize="11" level="3" pageId="35" pageNumber="36" reason="6">
|
||
<emphasis bold="true" box="[498,695,1444,1472]" italics="true" pageId="35" pageNumber="36">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="35.[498,1542,1444,1913]" pageId="35" pageNumber="36">In the seventh growth stage, the orbital notch in the frontal bone is pinched between the joint surfaces for the lacrimal and postorbital; the dorsal joint surface for the quadrate of the quadratojugal is wide and coarse, its rostral margin is concave, and its caudal half is developed into a wide ridge; the frontoparietal junction is wider than the sagittal crest rostral and caudal to it; and the joint surface for the dentary on the splenial is reinforced by a wide longitudinal ridge.</paragraph>
|
||
<paragraph blockId="35.[498,1542,1444,1913]" lastBlockId="36.[498,1542,234,340]" lastPageId="36" lastPageNumber="37" pageId="35" pageNumber="36">In the ninth growth stage, the joint surface for the quadratojugal on the jugal is positioned far caudal to the cornual process. In the tenth growth stage, the joint surface for the squamosal on the postorbital is coarsened by deep and numerous longitudinal ridges, the caudal shelf of the postorbital extends subtly ventrolaterally, and a ridge extends along the laterosphenoidoprootic suture. In the eleventh growth stage, the rostral margin of the parietal extends mediolaterally (i.e., it is not wedge-shaped), the postorbital buttress extends caudal to the midlength of the dorsotemporal fossa, and the lateral extent of the parietofrontal suture extends from mediolaterally to caudolaterally.</paragraph>
|
||
<paragraph blockId="36.[498,1542,373,642]" box="[498,809,373,401]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,809,373,401]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,809,373,401]" italics="true" pageId="36" pageNumber="37">Subcutaneous surface</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,373,642]" pageId="36" pageNumber="37">In the seventh growth stage, the subcutaneous surface of the ventral ramus of the lacrimal is coarse with papillae concentrated toward its rostral margin. In the eighth growth stage, lateral fossae are present on the alveolar process of the maxilla. In the ninth growth stage, the circumfossa ridge of the maxilla is low or absent, and the texture of the subcutaneous region of the chin is highly rugose. In the tenth growth stage, the groove that extends across the prefrontolacrimal process is deeply incised.</paragraph>
|
||
<paragraph blockId="36.[498,1542,674,784]" box="[498,828,674,702]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,828,674,702]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,828,674,702]" italics="true" pageId="36" pageNumber="37">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,674,784]" pageId="36" pageNumber="37">In the ninth growth stage, the cornual process of the postorbital exceeds the height of the bone.</paragraph>
|
||
<paragraph blockId="36.[498,1542,817,1166]" box="[498,822,817,845]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,822,817,845]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,822,817,845]" italics="true" pageId="36" pageNumber="37">Paranasal pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,817,1166]" pageId="36" pageNumber="37">In the eighth growth stage, the maxillary fenestra closely approaches the ventral margin of the external antorbital fenestra, the interfenestral strut is rostrocaudally narrow, the caudodorsal margin of the maxillary fenestra is straight or concave, the ventral margin of the maxillary fenestra is obliterated and incorporated into the antorbital fossa, and the mediolateral depth of the antorbital fossa is deep. In the ninth growth stage, the dorsal surface of the lacrimal extends dorsomedially, and the ventral edge of the subcutaneous surface at the septum ahead of the lacrimal pneumatic recess extends rostroventrally or grades into the antorbital fossa without a distinct orientation.</paragraph>
|
||
<paragraph blockId="36.[498,1542,1198,1348]" box="[498,857,1198,1226]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,857,1198,1226]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,857,1198,1226]" italics="true" pageId="36" pageNumber="37">Endoccipital pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,1198,1348]" pageId="36" pageNumber="37">In the eighth growth stage, the caudal surface of the paroccipital process is deeply concave and the caudodorsal surface is strongly convex. In the ninth growth stage, the bar that extends across the level of the lateral processes of the supraoccipital is prominent.</paragraph>
|
||
<paragraph blockId="36.[498,1542,1380,1610]" box="[498,839,1380,1408]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,839,1380,1408]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,839,1380,1408]" italics="true" pageId="36" pageNumber="37">Basicranial pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,1380,1610]" pageId="36" pageNumber="37">
|
||
In the eighth growth stage, the subcondylar fossa is obliterated by inflation. In the ninth growth stage, the pneumatic foramina of the subcondylar fossa are positioned far apart from each other (cf.
|
||
<bibRefCitation author="Witmer LM & Ridgley RC" box="[737,1015,1504,1530]" journalOrPublisher="Anatomical Record" pageId="36" pageNumber="37" pagination="1266 - 1296" part="292" refId="ref56657" refString="Witmer LM, Ridgley RC. 2009. New insights into the brain, braincase and ear region of tyrannosaurs (Dinosauria, Theropoda) with implications for sensory organization and behavior. Anatomical Record 292 (9): 1266 - 1296 DOI 10.1002 / ar. 20983." title="New insights into the brain, braincase and ear region of tyrannosaurs (Dinosauria, Theropoda) with implications for sensory organization and behavior" type="journal article" year="2009">
|
||
<emphasis box="[737,1015,1504,1530]" italics="true" pageId="36" pageNumber="37">Witmer & Ridgley, 2009</emphasis>
|
||
</bibRefCitation>
|
||
). In the eleventh growth stage, the rim of the caudoventral end of the subsellar recess is spout-like, and the ventral end of the preotic pendant is a low ridge.
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,1642,1793]" box="[498,787,1642,1670]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,787,1642,1670]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,787,1642,1670]" italics="true" pageId="36" pageNumber="37">Dorsotemporal fossa</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1542,1642,1793]" pageId="36" pageNumber="37">In the seventh growth stage, the sagittal crest of the frontal is equal to or greater than 37% the length of the bone, and the sagittal foramen is positioned far caudal to the rostral margin of the fossa (indicating rostralward encroachment of the musculature).</paragraph>
|
||
<paragraph blockId="36.[498,1541,1824,1934]" box="[498,681,1824,1852]" pageId="36" pageNumber="37">
|
||
<heading bold="true" box="[498,681,1824,1852]" fontSize="11" level="3" pageId="36" pageNumber="37" reason="6">
|
||
<emphasis bold="true" box="[498,681,1824,1852]" italics="true" pageId="36" pageNumber="37">Muscle scars</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="36.[498,1541,1824,1934]" lastBlockId="37.[498,1542,234,659]" lastPageId="37" lastPageNumber="38" pageId="36" pageNumber="37">In the seventh growth stage, the ridge along the squamosal process of the quadratojugal is mediolaterally wide in rostral view, and the nuchal crest is rugose but the texture does not reach the midline of the crest. In the eighth growth stage, the lateral half of the nuchal surface of the squamosal is coarsely textured, and the oval scar of the basisphenoid is oriented medioventrally. In the ninth growth stage, the oval scar faces lateroventrally and it is coarse in texture.</paragraph>
|
||
<paragraph blockId="37.[498,1542,234,659]" pageId="37" pageNumber="38">In the tenth growth stage, the rugose texture of the nuchal crest of the parietal reaches the midline of the crest, the ventral end of the metotic strut is scoured by the oval scar complex, and the basal tuber is coarse in texture. In the eleventh growth stage, the pits mediodorsal to the supraoccipital extend deeply into the nuchal crest, the dorsal rugosities of the nuchal crest extend onto the rostral surface of the crest, and the ventrolateral edge of the base of the rostrolateral process of the parietal is crossed by a subtle, ventrolaterally extending ridge.</paragraph>
|
||
<paragraph blockId="37.[498,1542,692,842]" box="[498,623,692,720]" pageId="37" pageNumber="38">
|
||
<heading bold="true" box="[498,623,692,720]" fontSize="11" level="3" pageId="37" pageNumber="38" reason="6">
|
||
<emphasis bold="true" box="[498,623,692,720]" italics="true" pageId="37" pageNumber="38">Dentition</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="37.[498,1542,692,842]" pageId="37" pageNumber="38">In the eighth growth stage, the first mesial alveolus of the dentary is substantially smaller than the alveoli at the middle of the tooth row (i.e., the second alveolus is enlarged), and the number of dentary alveoli is 13.</paragraph>
|
||
<paragraph blockId="37.[498,1542,876,1225]" box="[498,808,876,904]" pageId="37" pageNumber="38">
|
||
<heading bold="true" box="[498,808,876,904]" fontSize="11" level="3" pageId="37" pageNumber="38" reason="6">
|
||
<emphasis bold="true" box="[498,808,876,904]" italics="true" pageId="37" pageNumber="38">Appendicular skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="37.[498,1542,876,1225]" pageId="37" pageNumber="38">In the seventh growth stage, the fibula is stout, the proximal end of the bone is differentiated into distinct anterior and posterior knob-like convexities that produces a deeply concave dorsal margin, the anterior surface of the fibula distinctly widens toward its distal (=ventral) end, the medial fossa of the fibula is sharply differentiated into anterior and posterior fossae, the posterior extent of the medial fossa is a wide ridge that separates the fossa from the posterior edge of the bone, and the anterior surface of the bone above the iliofibularis tubercle (= bipartite scar) is wide and flat before becoming dorsally a narrow but blunt ridge.</paragraph>
|
||
<paragraph blockId="37.[498,1542,1264,1935]" box="[498,596,1264,1294]" pageId="37" pageNumber="38">
|
||
<heading bold="true" box="[498,596,1264,1294]" fontSize="12" level="2" pageId="37" pageNumber="38" reason="0">
|
||
<emphasis bold="true" box="[498,596,1264,1294]" pageId="37" pageNumber="38">Adults</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="37.[498,1542,1264,1935]" pageId="37" pageNumber="38">
|
||
Adults (=young adult of
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[792,1027,1310,1336]" journalOrPublisher="Nature" pageId="37" pageNumber="38" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[792,1027,1310,1336]" italics="true" pageId="37" pageNumber="38">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
) have a skull length of 1.3
|
||
<emphasis box="[1341,1356,1311,1336]" italics="true" pageId="37" pageNumber="38">–</emphasis>
|
||
1.4 m, a skull length to height ratio of 2.6
|
||
<emphasis box="[816,831,1351,1376]" italics="true" pageId="37" pageNumber="38">–</emphasis>
|
||
2.3, are greater than 22 years old, have an EFS, and, as is seen in subadults and young adults, initially have a high number of growth changes, which rapidly drops. Sixty-four synontomorphies occur in this category, and 13 of them have a CI of 1.0 (
|
||
<tableCitation box="[578,663,1470,1496]" captionStart="Table 9" captionStartId="38.[525,580,248,269]" captionTargetBox="[516,1524,407,959]" captionText="Table 9 Diagnostic characters of the adult and senescent growth categories of Tyrannosaurus rex. These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. The last row corresponds to the senescent adult character. PHYLO, corresponding homologous phylogenetic character (see Data S1)." httpUri="http://table.plazi.org/id/DF5C84E2FFE2FF8EFDEBFF7034F5FE3F" pageId="37" pageNumber="38" tableUuid="DF5C84E2FFE2FF8EFDEBFF7034F5FE3F">Table 9</tableCitation>
|
||
). In the skull, the greatest number of changes, 15 and 13, occurs in the snout and skull roof modules, respectively, and changes are seen in all modules (
|
||
<tableCitation box="[1380,1467,1509,1536]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="37" pageNumber="38" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
). Eight changes occur in the postcranium, where five occur in the pelvic girdle and limb, and three are seen in the axial skeleton. The adult stage corresponds to growth stages 13
|
||
<emphasis box="[1468,1483,1590,1615]" italics="true" pageId="37" pageNumber="38">–</emphasis>
|
||
20 of the ontogram (
|
||
<figureCitation box="[677,742,1629,1655]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="37" pageNumber="38">Fig. 2</figureCitation>
|
||
) and to the first part of the stationary phase of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1355,1424,1630,1655]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1355,1424,1630,1655]" italics="true" pageId="37" pageNumber="38">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth curve (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[580,813,1669,1695]" journalOrPublisher="Nature" pageId="37" pageNumber="38" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[580,813,1669,1695]" italics="true" pageId="37" pageNumber="38">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[828,907,1669,1695]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="37" pageNumber="38">Fig. 12</figureCitation>
|
||
). An ontogenetic progression among adults, based on size, was first hypothesized by
|
||
<bibRefCitation author="Paul GS" box="[861,996,1709,1735]" journalOrPublisher="New York: Simon and Schuster" pageId="37" pageNumber="38" refId="ref55555" refString="Paul GS. 1988. Predatory dinosaurs of the world: a complete illustrated guide. New York: Simon and Schuster." title="Predatory dinosaurs of the world: a complete illustrated guide" type="book" year="1988">
|
||
<emphasis box="[861,996,1709,1735]" italics="true" pageId="37" pageNumber="38">Paul (1988)</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<emphasis box="[1012,1023,1709,1735]" italics="true" pageId="37" pageNumber="38">“</emphasis>
|
||
AMNH FARB 50270 and other big
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1446,1515,1710,1735]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1446,1515,1710,1735]" italics="true" pageId="37" pageNumber="38">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens may or may not be subadults. This is possible because the biggest specimen (
|
||
<materialsCitation box="[506,686,1789,1815]" collectionCode="UCMP" pageId="37" pageNumber="38" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
)
|
||
<emphasis box="[694,722,1789,1814]" italics="true" pageId="37" pageNumber="38">…</emphasis>
|
||
is 29 percent longer than 5027
|
||
<emphasis box="[1082,1094,1788,1814]" italics="true" pageId="37" pageNumber="38">”</emphasis>
|
||
(1988: 344
|
||
<emphasis box="[1225,1240,1789,1814]" italics="true" pageId="37" pageNumber="38">–</emphasis>
|
||
345). Indeed, the results here recover this exact hypothesis of relative maturity (
|
||
<figureCitation box="[1150,1226,1828,1854]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="37" pageNumber="38">Figs. 1</figureCitation>
|
||
and
|
||
<figureCitation box="[1287,1301,1828,1854]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="37" pageNumber="38">2</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="37.[498,1542,1264,1935]" lastBlockId="38.[498,1542,1057,1443]" lastPageId="38" lastPageNumber="39" pageId="37" pageNumber="38">
|
||
Several mass estimates are available for adults (
|
||
<tableCitation box="[1069,1154,1868,1895]" captionStart="Table 5" captionStartId="25.[117,172,248,269]" captionTargetBox="[108,1524,407,885]" captionText="Table 5 Summary of mass estimates for Tyrannosaurus rex. Summary of published mass estimates (kg) for individual specimens of Tyrannosaurus rex with rank order in each column in parentheses; sum rank order given in right-hand column. The rank of AMNH FARB 5027 could not be identified given its presence in only one analysis and the great difference in estimates between the analyses. Column numbers: 1, Erickson et al. (2004); 2, Henderson & Snively (2004); 3, Bates et al. (2009); 4, Hutchinson et al. (2011); 5, Campione et al. (2014); 6, Snively et al. (2019); 7, Persons, Currie & Erickson (2019)." httpUri="http://table.plazi.org/id/DF5C84E2FFDDFFB1FF93FF703676FE3F" pageId="37" pageNumber="38" tableUuid="DF5C84E2FFDDFFB1FF93FF703676FE3F">Table 5</tableCitation>
|
||
). The mass for adults is 10,200 kg in
|
||
<bibRefCitation author="Henderson DM & Snively E." box="[530,868,1908,1934]" journalOrPublisher="Proceedings of the Royal Society B" pageId="37" pageNumber="38" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">Henderson & Snively (2004)</bibRefCitation>
|
||
, 6,071.8 kg in
|
||
<bibRefCitation author="Bates KT & Manning PL & Hodgetts D & Sellers WI" box="[1041,1251,1908,1934]" journalOrPublisher="PLOS ONE" pageId="37" pageNumber="38" pagination="e 4532" part="4" refId="ref52064" refString="Bates KT, Manning PL, Hodgetts D, Sellers WI. 2009. Estimating mass properties of dinosaurs using later imaging and 3 D computer modeling. PLOS ONE 4 (2): e 4532 DOI 10.1371 / journal. pone. 0004532." title="Estimating mass properties of dinosaurs using later imaging and 3 D computer modeling" type="journal article" year="2009">
|
||
<emphasis box="[1041,1251,1908,1934]" italics="true" pageId="37" pageNumber="38">Bates et al. (2009)</emphasis>
|
||
</bibRefCitation>
|
||
, 5,777
|
||
<emphasis box="[1328,1343,1909,1934]" italics="true" pageId="37" pageNumber="38">–</emphasis>
|
||
10,768 kg in
|
||
<bibRefCitation author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[498,779,1057,1084]" journalOrPublisher="PLOS ONE" pageId="38" pageNumber="39" pagination="e 97055" part="9" refId="ref54676" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLOS ONE 9 (5): e 97055 DOI 10.1371 / journal. pone. 0097055." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">
|
||
<emphasis box="[498,779,1057,1084]" italics="true" pageId="38" pageNumber="39">Hutchinson et al. (2011)</emphasis>
|
||
</bibRefCitation>
|
||
, 4,660
|
||
<emphasis box="[856,871,1058,1083]" italics="true" pageId="38" pageNumber="39">–</emphasis>
|
||
10,007 kg in
|
||
<bibRefCitation author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[1023,1288,1057,1084]" journalOrPublisher="Methods in Ecology and Evolution" pageId="38" pageNumber="39" pagination="913 - 923" part="5" refId="ref52505" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods in Ecology and Evolution 5 (9): 913 - 923 DOI 10.1111 / 2041 - 210 X. 12226." title="Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">
|
||
<emphasis box="[1023,1288,1057,1084]" italics="true" pageId="38" pageNumber="39">Campione et al. (2014)</emphasis>
|
||
</bibRefCitation>
|
||
, 6,986.6 kg in
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" journalOrPublisher="PeerJ" pageId="38" pageNumber="39" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="38" pageNumber="39">Snively et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
, and 5,112
|
||
<emphasis box="[768,783,1098,1123]" italics="true" pageId="38" pageNumber="39">–</emphasis>
|
||
8,870 kg in
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[923,1320,1097,1124]" journalOrPublisher="Anatomical Record" pageId="38" pageNumber="39" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[923,1320,1097,1124]" italics="true" pageId="38" pageNumber="39">Persons, Currie & Erickson (2019)</emphasis>
|
||
</bibRefCitation>
|
||
. The minimum estimated masses of adults are greater than 4,500 kg (
|
||
<bibRefCitation author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[1132,1402,1137,1163]" journalOrPublisher="PLOS ONE" pageId="38" pageNumber="39" pagination="e 97055" part="9" refId="ref54676" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLOS ONE 9 (5): e 97055 DOI 10.1371 / journal. pone. 0097055." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">
|
||
<emphasis box="[1132,1402,1137,1163]" italics="true" pageId="38" pageNumber="39">Hutchinson et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Campione NE & Evans DC & Brown CM & Carrano MT" journalOrPublisher="Methods in Ecology and Evolution" pageId="38" pageNumber="39" pagination="913 - 923" part="5" refId="ref52505" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods in Ecology and Evolution 5 (9): 913 - 923 DOI 10.1111 / 2041 - 210 X. 12226." title="Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">
|
||
<emphasis italics="true" pageId="38" pageNumber="39">Campione et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[641,1026,1177,1204]" journalOrPublisher="Anatomical Record" pageId="38" pageNumber="39" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[641,1026,1177,1204]" italics="true" pageId="38" pageNumber="39">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
), which overlaps with that of young adults (
|
||
<tableCitation box="[587,675,1217,1244]" captionStart="Table 5" captionStartId="25.[117,172,248,269]" captionTargetBox="[108,1524,407,885]" captionText="Table 5 Summary of mass estimates for Tyrannosaurus rex. Summary of published mass estimates (kg) for individual specimens of Tyrannosaurus rex with rank order in each column in parentheses; sum rank order given in right-hand column. The rank of AMNH FARB 5027 could not be identified given its presence in only one analysis and the great difference in estimates between the analyses. Column numbers: 1, Erickson et al. (2004); 2, Henderson & Snively (2004); 3, Bates et al. (2009); 4, Hutchinson et al. (2011); 5, Campione et al. (2014); 6, Snively et al. (2019); 7, Persons, Currie & Erickson (2019)." httpUri="http://table.plazi.org/id/DF5C84E2FFDDFFB1FF93FF703676FE3F" pageId="38" pageNumber="39" tableUuid="DF5C84E2FFDDFFB1FF93FF703676FE3F">Table 5</tableCitation>
|
||
). In contrast, the mean mass estimates for adults exceed those for young adults, indicating that a mass greater than 5,100 kg might be diagnostic for adults, whereas the range of mass in young adults is from ~3,000 kg to ~5,100 kg. Given the variety of techniques used for estimating mass in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[957,1024,1337,1362]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[957,1024,1337,1362]" italics="true" pageId="38" pageNumber="39">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, the differences in the results between them, and the incomplete sampling of specimens, mass is here only tentatively considered informative in distinguishing between the young adult and adult categories.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFE2FF8EFDEBFF7034F5FE3F" httpUri="http://table.plazi.org/id/DF5C84E2FFE2FF8EFDEBFF7034F5FE3F" pageId="38" pageNumber="39" startId="38.[525,580,248,269]" targetBox="[516,1524,407,959]" targetIsTable="true" targetPageId="38">
|
||
<paragraph blockId="38.[525,1515,248,387]" pageId="38" pageNumber="39">
|
||
<emphasis bold="true" box="[525,1515,248,270]" pageId="38" pageNumber="39">
|
||
Table 9 Diagnostic characters of the adult and senescent growth categories of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1322,1509,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1322,1509,249,270]" italics="true" pageId="38" pageNumber="39">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. The last row corresponds to the senescent adult character. PHYLO, corresponding homologous phylogenetic character (see Data S1).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="38" pageNumber="39">
|
||
<table box="[516,1524,407,959]" gridcols="2" gridrows="15" pageId="38" pageNumber="39">
|
||
<tr box="[516,1524,407,429]" gridrow="0" pageId="38" pageNumber="39">
|
||
<th box="[516,1287,407,429]" gridcol="0" gridrow="0" pageId="38" pageNumber="39">Character</th>
|
||
<th box="[1372,1524,407,429]" gridcol="1" gridrow="0" pageId="38" pageNumber="39">
|
||
<emphasis bold="true" box="[1372,1524,407,429]" pageId="38" pageNumber="39">Growth change</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,449,471]" gridrow="1" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,449,471]" gridcol="0" gridrow="1" pageId="38" pageNumber="39">147. Cervical vertebrae, centrum, hypapophysis, presence (PHYLO 280)</td>
|
||
<td box="[1372,1524,449,471]" gridcol="1" gridrow="1" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,486,509]" gridrow="2" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,486,509]" gridcol="0" gridrow="2" pageId="38" pageNumber="39">166. Ilium, pubic peduncle, ventral margin, orientation (PHYLO 333)</td>
|
||
<td box="[1372,1524,486,509]" gridcol="1" gridrow="2" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,524,546]" gridrow="3" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,524,546]" gridcol="0" gridrow="3" pageId="38" pageNumber="39">546. Jugal, medial maxillary process, rostral end, depth inset</td>
|
||
<td box="[1372,1524,524,546]" gridcol="1" gridrow="3" pageId="38" pageNumber="39">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,561,583]" gridrow="4" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,561,583]" gridcol="0" gridrow="4" pageId="38" pageNumber="39">590. Jugal, ventral quadratojugal process, dorsolateral surface</td>
|
||
<td box="[1372,1524,561,583]" gridcol="1" gridrow="4" pageId="38" pageNumber="39">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,599,621]" gridrow="5" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,599,621]" gridcol="0" gridrow="5" pageId="38" pageNumber="39">671. Squamosal, quadratojugal process, ventral margin, concavity</td>
|
||
<td box="[1372,1524,599,621]" gridcol="1" gridrow="5" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,637,659]" gridrow="6" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,637,659]" gridcol="0" gridrow="6" pageId="38" pageNumber="39">694. Squamosal, joint surface for otoccipital, form</td>
|
||
<td box="[1372,1524,637,659]" gridcol="1" gridrow="6" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,674,696]" gridrow="7" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,674,696]" gridcol="0" gridrow="7" pageId="38" pageNumber="39">735. Quadratojugal, jugal ramus, ventral margin, form</td>
|
||
<td box="[1372,1524,674,696]" gridcol="1" gridrow="7" pageId="38" pageNumber="39">1 ==> 2</td>
|
||
</tr>
|
||
<tr box="[516,1524,712,734]" gridrow="8" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,712,734]" gridcol="0" gridrow="8" pageId="38" pageNumber="39">1051. Basisphenoid, lateral margin, form</td>
|
||
<td box="[1372,1524,712,734]" gridcol="1" gridrow="8" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,750,772]" gridrow="9" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,750,772]" gridcol="0" gridrow="9" pageId="38" pageNumber="39">1223. Axis, axial intercentrum, joint surface, ratio height to mediolateral width</td>
|
||
<td box="[1372,1524,750,772]" gridcol="1" gridrow="9" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,787,809]" gridrow="10" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,787,809]" gridcol="0" gridrow="10" pageId="38" pageNumber="39">1256. Axis, spinous process, dorsolateral process, dorsoventral height</td>
|
||
<td box="[1372,1524,787,809]" gridcol="1" gridrow="10" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,825,847]" gridrow="11" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,825,847]" gridcol="0" gridrow="11" pageId="38" pageNumber="39">1613. Ischium, semicircular scars, number</td>
|
||
<td box="[1372,1524,825,847]" gridcol="1" gridrow="11" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,862,884]" gridrow="12" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,862,884]" gridcol="0" gridrow="12" pageId="38" pageNumber="39">1668. Fibula, distal end, anterior margin, form</td>
|
||
<td box="[1372,1524,862,884]" gridcol="1" gridrow="12" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,900,921]" gridrow="13" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,900,921]" gridcol="0" gridrow="13" pageId="38" pageNumber="39">1849. EFS, presence</td>
|
||
<td box="[1372,1524,900,921]" gridcol="1" gridrow="13" pageId="38" pageNumber="39">0 ==> 1</td>
|
||
</tr>
|
||
<tr box="[516,1524,937,959]" gridrow="14" pageId="38" pageNumber="39">
|
||
<td box="[516,1287,937,959]" gridcol="0" gridrow="14" pageId="38" pageNumber="39">1220. Axis, axial intercentrum, anterior joint surface, ventral margin, separation</td>
|
||
<td box="[1372,1524,937,959]" gridcol="1" gridrow="14" pageId="38" pageNumber="39">1 ==> 2</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="38.[498,1542,1481,1910]" box="[498,741,1481,1509]" pageId="38" pageNumber="39">
|
||
<heading bold="true" box="[498,741,1481,1509]" fontSize="11" level="3" pageId="38" pageNumber="39" reason="6">
|
||
<emphasis bold="true" box="[498,741,1481,1509]" italics="true" pageId="38" pageNumber="39">Skull architecture</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="38.[498,1542,1481,1910]" pageId="38" pageNumber="39">In the thirteenth growth stage, the lateral depression of the jugal is shallow and enhanced by ridges, the surangular shelf extends lateroventrally, the angulation in the ventral margin of the maxilla is distinct, the ventral jugal process of the maxilla is a massive convex strut, the rostrodorsal margin of the rostral ramus of the lacrimal extends sharply rostroventrally, the rostral end of the medial maxillary process of the jugal above the joint surface for the maxilla is sharply inset caudally, the base of the medial process of the squamosal is separated by a notch from the margin of the dorsotemporal fenestra, the caudodorsomedial edge of the shaft of the quadrate is distinctly concave, the rostral margin of the vomeropterygoid process of the palatine is positioned ahead of the rostral palatine recess, and the lateral margin of the basisphenoid is deeply embayed.</paragraph>
|
||
<paragraph blockId="39.[498,1542,234,1177]" pageId="39" pageNumber="40">In the fourteenth growth stage, the narrow caudal end of the ascending ramus of the maxilla above the internal antorbital fenestra twists along its course, the tip of the rostrodorsal process of the lacrimal extends rostrally past the midlength of the internal antorbital fenestra, the lateral depression of the postorbital process of the jugal reaches the level of the rostral end of the postorbital contact, the parietal is elaborated into a ridge above each of the tines of the dorsal process of the supraoccipital, the Meckelian groove of the dentary is positioned at the midheight of the bone rostrally and above the midheight caudally, and the caudal margin of the dentary is deeply concave.</paragraph>
|
||
<paragraph blockId="39.[498,1542,234,1177]" pageId="39" pageNumber="40">In the fifteenth growth stage, the convex dorsal margin of the dentary extends to alveolus 7. In the seventeenth growth stage, the dorsal margin of the postorbital is swollen such that the medial eversion is completely obscured. In the eighteenth growth stage, the medial edge of the caudal margin of the quadratojugal is positioned caudal to the lateral edge of the bone. In the nineteenth growth stage, the rostralmost incursion of the internal antorbital fenestra is at the level of the fifth tooth from the caudal end of the tooth row, the rostroventral margin of the ventral ramus of the lacrimal is straight, the lateral ridge of the ventral postorbital process of the squamosal extends more ventrally than laterally, the dorsal margin of the jugal ramus of the quadratojugal extends to the medial edge of the shaft, and the ventral margin of the ramus if ventrally convex.</paragraph>
|
||
<paragraph blockId="39.[498,1542,234,1177]" pageId="39" pageNumber="40">In the twentieth growth stage, the quadrate fossa of the quadratojugal is shallow in depth, the ventral ramus of the lacrimal below the medial pneumatic recess is convex in vertical section, the nasal ramus of the frontal is wide and truncated, a ventrally-extending flange is absent from the ventral margin of the basioccipital, and the Meckelian groove is positioned below the midheight of the dentary rostrally and above midheight caudally.</paragraph>
|
||
<paragraph blockId="39.[498,1542,1216,1884]" box="[498,695,1216,1244]" pageId="39" pageNumber="40">
|
||
<heading bold="true" box="[498,695,1216,1244]" fontSize="11" level="3" pageId="39" pageNumber="40" reason="6">
|
||
<emphasis bold="true" box="[498,695,1216,1244]" italics="true" pageId="39" pageNumber="40">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="39.[498,1542,1216,1884]" pageId="39" pageNumber="40">In the thirteenth growth stage, the dorsolateral surface of the ventral quadratojugal process of the jugal is dominated by a single large ridge, the dorsal margin of the joint surface for the quadrate of the ventral quadrate process of the quadratojugal extends rostrodorsally, the joint surface for the pterygoid on the caudal process of the ectopterygoid is deeply excavated, and the joint surface for the splenial on the ventral bar of the dentary has a peg-in-socket form.</paragraph>
|
||
<paragraph blockId="39.[498,1542,1216,1884]" pageId="39" pageNumber="40">In the fourteenth growth stage, the condylar surface of the occipital condyle has a distinct marginal rim. In the sixteenth growth stage, the joint surface for the splenial on the lingual bar of the dentary is inset. In the eighteenth growth stage, the concavity in the ventral margin of the quadratojugal process of the squamosal is absent, and the joint surface for the otoccipital on the squamosal is convex.</paragraph>
|
||
<paragraph blockId="39.[498,1542,1216,1884]" lastBlockId="40.[498,1541,234,301]" lastPageId="40" lastPageNumber="41" pageId="39" pageNumber="40">In the nineteenth growth stage, the distal end of the jugal ramus of the postorbital is wedge-shaped to fit into a complementary groove in the jugal, the joint surface for the medial frontal process of the nasal is long and reaches caudally past the rostral end of the joint surface for the prefrontal, the rostral margin of the Meckelian fossa extends ventrolaterally to the medial surface of the bone, and the prearticuloangular buttress is present. In the twentieth growth stage, the joint surface for the palatine is present medially and rostrally on the rostroventral ala of the lacrimal.</paragraph>
|
||
<paragraph blockId="40.[498,1540,332,482]" box="[498,822,332,360]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,822,332,360]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,822,332,360]" italics="true" pageId="40" pageNumber="41">Paranasal pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1540,332,482]" pageId="40" pageNumber="41">In the thirteenth growth stage, the distal recess of the rostral ramus of the lacrimal faces more ventrally than laterally. In the nineteenth growth stage, a strut extends across the medial wall of the secondary fossa of the jugal.</paragraph>
|
||
<paragraph blockId="40.[498,1541,514,623]" box="[498,839,514,542]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,839,514,542]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,839,514,542]" italics="true" pageId="40" pageNumber="41">Basicranial pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1541,514,623]" pageId="40" pageNumber="41">In the twentieth growth stage, the subcondylar foramen of the basioccipital emits a deeply inset channel from its lower margin.</paragraph>
|
||
<paragraph blockId="40.[498,1542,655,964]" box="[498,741,655,683]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,741,655,683]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,741,655,683]" italics="true" pageId="40" pageNumber="41">Neurovasculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1542,655,964]" pageId="40" pageNumber="41">In the thirteenth growth stage, the medial surface of the ventral ramus of the lacrimal is scoured by deep sulci. In the fourteenth growth stage, the subnarial foramen produces a deep notch in the rostral margin of the maxilla, and lateromedially penetrating foramina in the antorbital fossa of the maxilla are present. In the eighteenth growth stage, the caudal surface of the jugal process of the ectopterygoid is not perforated by a foramen, and lateromedially penetrating foramina are absent from the antorbital fossa of the maxilla.</paragraph>
|
||
<paragraph blockId="40.[498,1541,996,1106]" box="[498,787,996,1024]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,787,996,1024]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,787,996,1024]" italics="true" pageId="40" pageNumber="41">Dorsotemporal fossa</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1541,996,1106]" pageId="40" pageNumber="41">In the thirteenth growth stage, the rostral margin of the dorsotemporal fossa of the frontal is elaborated into a ridge or crest.</paragraph>
|
||
<paragraph blockId="40.[498,1485,1138,1248]" box="[498,809,1138,1166]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,809,1138,1166]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,809,1138,1166]" italics="true" pageId="40" pageNumber="41">Subcutaneous surface</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1485,1138,1248]" pageId="40" pageNumber="41">In the thirteenth growth stage, the texture of the caudal process of the squamosal is coarsely rugose.</paragraph>
|
||
<paragraph blockId="40.[498,1464,1280,1350]" box="[498,828,1280,1308]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,828,1280,1308]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,828,1280,1308]" italics="true" pageId="40" pageNumber="41">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1464,1280,1350]" box="[498,1464,1323,1350]" pageId="40" pageNumber="41">In the nineteenth growth stage, the epipostorbital is fused to the underlying bone.</paragraph>
|
||
<paragraph blockId="40.[498,1542,1382,1532]" box="[498,623,1382,1410]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,623,1382,1410]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,623,1382,1410]" italics="true" pageId="40" pageNumber="41">Dentition</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1542,1382,1532]" pageId="40" pageNumber="41">In the thirteenth growth stage, the number of dentary teeth is 14. In the seventeenth growth stage, the rostral margins of the interdental plates are notched (almost certainly an epiphenomenon of tooth enlargement).</paragraph>
|
||
<paragraph blockId="40.[498,1541,1563,1713]" box="[498,692,1563,1591]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,692,1563,1591]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,692,1563,1591]" italics="true" pageId="40" pageNumber="41">Axial skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1541,1563,1713]" pageId="40" pageNumber="41">In the fifteenth growth stage, the joint surface of the axial intercentrum is dorsoventrally deep, and the spinous process of the axis is dorsoventrally massive. In the nineteenth growth stage, the hypophysis is present on the cervical centra.</paragraph>
|
||
<paragraph blockId="40.[498,1542,1745,1934]" box="[498,808,1745,1773]" pageId="40" pageNumber="41">
|
||
<heading bold="true" box="[498,808,1745,1773]" fontSize="11" level="3" pageId="40" pageNumber="41" reason="6">
|
||
<emphasis bold="true" box="[498,808,1745,1773]" italics="true" pageId="40" pageNumber="41">Appendicular skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="40.[498,1542,1745,1934]" lastBlockId="41.[498,1542,234,300]" lastPageId="41" lastPageNumber="42" pageId="40" pageNumber="41">In the fourteenth growth stage, the ischium has two semicircular scars. In the eighteenth growth stage, the ventral margin of the public peduncle of the ilium extends horizontally and the anterior surface of the fibula below the bipartite scar is narrow. In the nineteenth growth stage, the distal end of the shaft of the fibula flares abruptly to the ventral joint surface that produces a deep crease in the shaft, and the anterior margin of the distal end of the fibula bulges anteriorly.</paragraph>
|
||
<paragraph blockId="41.[498,1542,343,894]" box="[498,759,343,373]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,759,343,373]" fontSize="12" level="2" pageId="41" pageNumber="42" reason="0">
|
||
<emphasis bold="true" box="[498,759,343,373]" pageId="41" pageNumber="42">Senescent adults</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1542,343,894]" pageId="41" pageNumber="42">
|
||
Finally, senescent adults have a skull length of at least 1.4 m, are at least 28 years old, and have an EFS. In the skull, the greatest number of changes occurs in the skull roof module, and the rest of the synontomorphies are limited to the snout, suspensorium, and the braincase modules, and postcranial changes occur in the axis (
|
||
<tableCitation box="[1235,1323,508,535]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="41" pageNumber="42" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
). Of the 19 synontomorphies that support this growth stage, only one has a CI of 1.0 (
|
||
<tableCitation box="[1383,1471,548,575]" captionStart="Table 9" captionStartId="38.[525,580,248,269]" captionTargetBox="[516,1524,407,959]" captionText="Table 9 Diagnostic characters of the adult and senescent growth categories of Tyrannosaurus rex. These characters are unambiguously optimized on the topology and have a consistency index (CI) of 1.0 indicating that they have changed only once on the ontogram and so can be considered diagnostic of the growth category. The last row corresponds to the senescent adult character. PHYLO, corresponding homologous phylogenetic character (see Data S1)." httpUri="http://table.plazi.org/id/DF5C84E2FFE2FF8EFDEBFF7034F5FE3F" pageId="41" pageNumber="42" tableUuid="DF5C84E2FFE2FF8EFDEBFF7034F5FE3F">Table 9</tableCitation>
|
||
). This category corresponds to growth stage 21 of the ontogram (
|
||
<figureCitation box="[1226,1289,588,615]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="41" pageNumber="42">Fig. 2</figureCitation>
|
||
) and to the end of the stationary phase of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[770,837,629,654]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[770,837,629,654]" italics="true" pageId="41" pageNumber="42">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth curve (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1016,1247,628,654]" journalOrPublisher="Nature" pageId="41" pageNumber="42" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1016,1247,628,654]" italics="true" pageId="41" pageNumber="42">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1260,1338,628,654]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="41" pageNumber="42">Fig. 12</figureCitation>
|
||
); there is a single exemplar specimen,
|
||
<materialsCitation box="[739,926,668,695]" collectionCode="FMNH" pageId="41" pageNumber="42" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1542,343,894]" pageId="41" pageNumber="42">
|
||
The estimated mass for FMNH PR208 is greater than those obtained for other adults in
|
||
<bibRefCitation author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[530,813,748,774]" journalOrPublisher="PLOS ONE" pageId="41" pageNumber="42" pagination="e 97055" part="9" refId="ref54676" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLOS ONE 9 (5): e 97055 DOI 10.1371 / journal. pone. 0097055." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">
|
||
<emphasis box="[530,813,748,774]" italics="true" pageId="41" pageNumber="42">Hutchinson et al. (2011)</emphasis>
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[874,1104,748,774]" journalOrPublisher="PeerJ" pageId="41" pageNumber="42" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[874,1104,748,774]" italics="true" pageId="41" pageNumber="42">Snively et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
(
|
||
<tableCitation box="[1123,1210,748,775]" captionStart="Table 5" captionStartId="25.[117,172,248,269]" captionTargetBox="[108,1524,407,885]" captionText="Table 5 Summary of mass estimates for Tyrannosaurus rex. Summary of published mass estimates (kg) for individual specimens of Tyrannosaurus rex with rank order in each column in parentheses; sum rank order given in right-hand column. The rank of AMNH FARB 5027 could not be identified given its presence in only one analysis and the great difference in estimates between the analyses. Column numbers: 1, Erickson et al. (2004); 2, Henderson & Snively (2004); 3, Bates et al. (2009); 4, Hutchinson et al. (2011); 5, Campione et al. (2014); 6, Snively et al. (2019); 7, Persons, Currie & Erickson (2019)." httpUri="http://table.plazi.org/id/DF5C84E2FFDDFFB1FF93FF703676FE3F" pageId="41" pageNumber="42" tableUuid="DF5C84E2FFDDFFB1FF93FF703676FE3F">Table 5</tableCitation>
|
||
). However, its mass is exceeded by an adult in the comprehensive samples of
|
||
<bibRefCitation author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[1131,1391,788,814]" journalOrPublisher="Methods in Ecology and Evolution" pageId="41" pageNumber="42" pagination="913 - 923" part="5" refId="ref52505" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods in Ecology and Evolution 5 (9): 913 - 923 DOI 10.1111 / 2041 - 210 X. 12226." title="Body mass estimation in on-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">
|
||
<emphasis box="[1131,1391,788,814]" italics="true" pageId="41" pageNumber="42">Campione et al. (2014)</emphasis>
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" journalOrPublisher="Anatomical Record" pageId="41" pageNumber="42" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="41" pageNumber="42">Persons, Currie & Erickson (2019)</emphasis>
|
||
</bibRefCitation>
|
||
; therefore, the mass of senescent adults does not exceed the range seen in adults and so it cannot be used to diagnose this growth stage.
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1542,930,1080]" box="[498,741,930,958]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,741,930,958]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,741,930,958]" italics="true" pageId="41" pageNumber="42">Skull architecture</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1542,930,1080]" pageId="41" pageNumber="42">The antorbital fossa below the internal antorbital fenestra is extremely shallow, low ridges extend into the quadrate cotyle of the squamosal, and the orbital notch is a deep cleft in the orbital surface of the frontal.</paragraph>
|
||
<paragraph blockId="41.[498,1541,1117,1307]" box="[498,695,1117,1145]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,695,1117,1145]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,695,1117,1145]" italics="true" pageId="41" pageNumber="42">Joint surfaces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1541,1117,1307]" pageId="41" pageNumber="42">The joint surface for the palatine on the rostroventral ala of the lacrimal is present medially only, the crease between the postorbital buttress and shelf of the frontal is present, and the joint surface for the prefrontal in the orbital surface of the frontal is positioned close to the orbital notch.</paragraph>
|
||
<paragraph blockId="41.[498,1080,1344,1414]" box="[498,809,1344,1372]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,809,1344,1372]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,809,1344,1372]" italics="true" pageId="41" pageNumber="42">Subcutaneous surface</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1080,1344,1414]" box="[498,1080,1388,1414]" pageId="41" pageNumber="42">The dorsolateral surface of the lacrimal is rugose.</paragraph>
|
||
<paragraph blockId="41.[498,1529,1451,1560]" box="[498,828,1451,1479]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,828,1451,1479]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,828,1451,1479]" italics="true" pageId="41" pageNumber="42">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1529,1451,1560]" pageId="41" pageNumber="42">The cornual process of the postorbital does not interrupt the rostrolateral corner of the dorsotemporal fossa.</paragraph>
|
||
<paragraph blockId="41.[498,1502,1598,1708]" box="[498,822,1598,1626]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,822,1598,1626]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,822,1598,1626]" italics="true" pageId="41" pageNumber="42">Paranasal pneumaticity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1502,1598,1708]" pageId="41" pageNumber="42">The caudalmost extent of the trough medial to the tooth root bulges is limited to the maxillary antrum.</paragraph>
|
||
<paragraph blockId="41.[498,1542,1745,1934]" box="[498,741,1745,1773]" pageId="41" pageNumber="42">
|
||
<heading bold="true" box="[498,741,1745,1773]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="6">
|
||
<emphasis bold="true" box="[498,741,1745,1773]" italics="true" pageId="41" pageNumber="42">Neurovasculature</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="41.[498,1542,1745,1934]" pageId="41" pageNumber="42">The medial surface of the ventral ramus of the lacrimal is smooth, the foramen that penetrates the orbital surface of the jugal ramus of the postorbital is far medial to the lateral margin of the bone, and the ventral surface of the basituberal web is not penetrated by a foramen.</paragraph>
|
||
<paragraph blockId="42.[498,1541,233,423]" box="[498,681,233,261]" pageId="42" pageNumber="43">
|
||
<heading bold="true" box="[498,681,233,261]" fontSize="11" level="3" pageId="42" pageNumber="43" reason="6">
|
||
<emphasis bold="true" box="[498,681,233,261]" italics="true" pageId="42" pageNumber="43">Muscle scars</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1541,233,423]" pageId="42" pageNumber="43">The suborbital ligament scar is convex or bulbous, the lateral surface between the jugal process and ventral quadrate process of the quadratojugal is scoured by a deep fossa that is secured by a ridge, and the mediolaterally extending ridge ahead of the parietofrontal suture is absent.</paragraph>
|
||
<paragraph blockId="42.[498,1542,461,730]" box="[498,692,461,489]" pageId="42" pageNumber="43">
|
||
<heading bold="true" box="[498,692,461,489]" fontSize="11" level="3" pageId="42" pageNumber="43" reason="6">
|
||
<emphasis bold="true" box="[498,692,461,489]" italics="true" pageId="42" pageNumber="43">Axial skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1542,461,730]" pageId="42" pageNumber="43">The anterior surface of the axial intercentrum is deeply concave and bounded ventrally by a prominent rim, a deep groove separates the ventral edge of the anterior joint surface of the intercentrum from the ventral surface of the bone, the anterior margin of the intercentrum is concave and the posterior margin is convex, and the concave ventral surface of the intercentrum is limited to the ventral and lateroventral surfaces of the bone by the swollen parapophysis.</paragraph>
|
||
<paragraph blockId="42.[498,1542,776,928]" box="[498,909,776,806]" pageId="42" pageNumber="43">
|
||
<heading bold="true" box="[498,909,776,806]" fontSize="12" level="2" pageId="42" pageNumber="43" reason="0">
|
||
<emphasis bold="true" box="[498,909,776,806]" pageId="42" pageNumber="43">Craniomandibular variation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1542,776,928]" pageId="42" pageNumber="43">
|
||
The results obtained here provide an opportunity to review the variation reported by
|
||
<bibRefCitation author="Molnar RE" box="[498,665,861,887]" journalOrPublisher="Palaeontographica Abteilung A" pageId="42" pageNumber="43" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[498,665,861,887]" italics="true" pageId="42" pageNumber="43">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
in his extensive descriptive monograph of the skull and mandibular rami of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,567,902,927]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="42" pageNumber="43" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,902,927]" italics="true" pageId="42" pageNumber="43">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and recast in the context of ontogeny.
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1542,966,1913]" box="[498,704,966,994]" pageId="42" pageNumber="43">
|
||
<heading bold="true" box="[498,704,966,994]" fontSize="11" level="3" pageId="42" pageNumber="43" reason="6">
|
||
<emphasis bold="true" box="[498,704,966,994]" italics="true" pageId="42" pageNumber="43">Facial skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1542,966,1913]" pageId="42" pageNumber="43">
|
||
The foramen in the base of the antorbital fossa of the maxilla in
|
||
<materialsCitation box="[1236,1393,1010,1036]" collectionCode="LACM" pageId="42" pageNumber="43" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
is individual variation. The difference in shape of the maxillary fenestra seen between adults is almost certainly an artifact of variability in the form of its caudodorsal margin, ventral margin, and rostral margin. Comparison of the shapes of the fenestra (trapezoidal, oval, triangular) described by
|
||
<bibRefCitation author="Molnar RE" box="[788,957,1169,1196]" journalOrPublisher="Palaeontographica Abteilung A" pageId="42" pageNumber="43" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[788,957,1169,1196]" italics="true" pageId="42" pageNumber="43">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
with the ontogram do not show a sequential pattern. The absence of
|
||
<emphasis box="[783,795,1209,1235]" italics="true" pageId="42" pageNumber="43">“</emphasis>
|
||
discernable sculpture
|
||
<emphasis box="[1042,1054,1209,1235]" italics="true" pageId="42" pageNumber="43">”</emphasis>
|
||
on the maxilla of
|
||
<materialsCitation box="[1272,1508,1209,1236]" collectionCode="TMP" pageId="42" pageNumber="43" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Molnar RE" box="[508,725,1249,1276]" journalOrPublisher="Palaeontographica Abteilung A" pageId="42" pageNumber="43" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[508,665,1249,1275]" italics="true" pageId="42" pageNumber="43">Molnar, 1990</emphasis>
|
||
: 142
|
||
</bibRefCitation>
|
||
), taken here to mean the absence of fossae on the horizontal ramus, represents individual variation.
|
||
</paragraph>
|
||
<paragraph blockId="42.[498,1542,966,1913]" pageId="42" pageNumber="43">
|
||
The difference in the angle and position of the flexure of the jugal ramus of the maxilla might correspond to relative maturity, where it changes from rostral and distinct (e.g.,
|
||
<materialsCitation box="[562,797,1408,1435]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="42" pageNumber="43" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[810,966,1409,1435]" collectionCode="SDSM" pageId="42" pageNumber="43" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
) to caudal and indistinct (e.g.,
|
||
<materialsCitation box="[1334,1498,1409,1435]" collectionCode="LACM" pageId="42" pageNumber="43" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
). Likewise, the change from a large third maxillary antrum chamber (e.g.,
|
||
<materialsCitation box="[1356,1517,1448,1474]" collectionCode="LACM" pageId="42" pageNumber="43" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
;
|
||
<bibRefCitation author="Molnar RE" box="[498,655,1488,1514]" journalOrPublisher="Palaeontographica Abteilung A" pageId="42" pageNumber="43" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[498,655,1488,1514]" italics="true" pageId="42" pageNumber="43">Molnar, 1990</emphasis>
|
||
</bibRefCitation>
|
||
:
|
||
<figureCitation box="[670,737,1488,1514]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="42" pageNumber="43">Fig. 2</figureCitation>
|
||
) to a large second chamber (e.g.,
|
||
<materialsCitation box="[1136,1314,1488,1514]" collectionCode="UCMP" pageId="42" pageNumber="43" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
) corresponds to an increase in relative maturity. The foramen in the caudodorsal surface of the interfenestral strut in
|
||
<materialsCitation box="[756,866,1568,1594]" collectionCode="CM" pageId="42" pageNumber="43" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
is individual variation. The dorsoventral position of the palatal process of the maxilla is broadly congruent with relative maturity, where it is ventral in position in relatively immature adults (e.g.,
|
||
<materialsCitation box="[1132,1366,1647,1674]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="42" pageNumber="43" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1379,1534,1648,1674]" collectionCode="SDSM" pageId="42" pageNumber="43" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
) and dorsal in relatively mature adults (
|
||
<materialsCitation box="[940,1120,1688,1714]" collectionCode="UCMP" pageId="42" pageNumber="43" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
). However, these conditions are seen in specimens that share growth stage 17 (ventral in
|
||
<materialsCitation box="[1111,1223,1727,1753]" collectionCode="CM" pageId="42" pageNumber="43" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
, dorsal in
|
||
<materialsCitation box="[1348,1512,1727,1753]" collectionCode="LACM" pageId="42" pageNumber="43" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
). This character was not included in the character matrix; it may turn out to separate the specimens at growth stage 17 from each other, if, and only if, the character states can be verified and the analysis re-run with them included. The absence of interdental pits from the maxilla are individual variation unique to
|
||
<materialsCitation box="[1043,1199,1887,1913]" collectionCode="SDSM" pageId="42" pageNumber="43" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,234,978]" pageId="43" pageNumber="44">
|
||
Although
|
||
<bibRefCitation author="Molnar RE" box="[649,817,234,260]" journalOrPublisher="Palaeontographica Abteilung A" pageId="43" pageNumber="44" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[649,817,234,260]" italics="true" pageId="43" pageNumber="44">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
regarded absence of sculpture on the subcutaneous surface of the maxilla, lacrimal, and nasal in
|
||
<materialsCitation box="[906,1142,274,301]" collectionCode="TMP" pageId="43" pageNumber="44" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
as evidence for immaturity, based on the results here it is individual variation, as well as the coarse texture of the nasal that is seen in
|
||
<materialsCitation box="[672,791,354,380]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
. The difference between flattened premaxillary processes of the nasal in
|
||
<materialsCitation box="[638,868,393,420]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
and the rod-like processes in
|
||
<materialsCitation box="[1213,1370,394,420]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
and
|
||
<materialsCitation box="[1428,1542,394,420]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
are consistent with increasing relative maturity.
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,234,978]" pageId="43" pageNumber="44">
|
||
<emphasis box="[530,715,473,499]" italics="true" pageId="43" pageNumber="44">Molnar’ s (1990)</emphasis>
|
||
comments on the relative development of the cornual process of the postorbital deserve some comment. The process is a composite structure that includes a rugosity that is an intrinsic part of the bone to which an overlying osteoderm, termed here the epipostorbital, is attached. The epipostorbital is sometimes difficult to distinguish from the underlying postorbital, but it is clearly distinguished by its distinctly flat ventral surface along which a distinct, tuberculate, and eave-like flange extends (
|
||
<bibRefCitation author="Molnar RE" box="[1359,1516,673,699]" journalOrPublisher="Palaeontographica Abteilung A" pageId="43" pageNumber="44" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[1359,1516,673,699]" italics="true" pageId="43" pageNumber="44">Molnar, 1990</emphasis>
|
||
</bibRefCitation>
|
||
). The epipostorbital is not preserved in
|
||
<materialsCitation box="[950,1112,712,738]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
, which accounts for its difference from the other specimens. The progression in development regarded by
|
||
<bibRefCitation author="Molnar RE" box="[1354,1523,752,778]" journalOrPublisher="Palaeontographica Abteilung A" pageId="43" pageNumber="44" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[1354,1523,752,778]" italics="true" pageId="43" pageNumber="44">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
, from
|
||
<materialsCitation box="[565,718,792,818]" collectionCode="SDSM" pageId="43" pageNumber="44" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
to
|
||
<materialsCitation box="[759,992,792,819]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
to
|
||
<materialsCitation box="[1033,1151,792,818]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
, does follow the progression of maturity found here. However, the reduced condition of the epipostorbital among the most mature specimens (growth stages 19, 20), indicates a reversal of the pattern. A growth-related reduction in ornamentation is seen in other dinosaurs (e.g.,
|
||
<taxonomicName baseAuthorityName="Horner & Goodwin" baseAuthorityYear="2006" box="[1406,1535,912,937]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="43" pageNumber="44" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1406,1535,912,937]" italics="true" pageId="43" pageNumber="44">Triceratops</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Horner & Goodwin" baseAuthorityYear="2009" box="[498,729,951,977]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="43" pageNumber="44" phylum="Chordata" rank="genus">
|
||
<emphasis box="[498,729,951,977]" italics="true" pageId="43" pageNumber="44">Pachycephalosaurus</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[744,1043,951,978]" journalOrPublisher="Proceedings of the Royal Society of London: Biology" pageId="43" pageNumber="44" pagination="2757 - 2761" part="273" refId="ref54486" refString="Horner JR, Goodwin MB. 2006. Major cranial changes during Triceratops ontogeny. Proceedings of the Royal Society of London: Biology 273 (1602): 2757 - 2761 DOI 10.1098 / rspb. 2006.3643." title="Major cranial changes during Triceratops ontogeny" type="journal article" year="2006">
|
||
<emphasis box="[744,1043,951,978]" italics="true" pageId="43" pageNumber="44">Horner & Goodwin, 2006</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[1058,1356,951,978]" journalOrPublisher="PLOS ONE" pageId="43" pageNumber="44" pagination="e 7626" part="4" refId="ref54525" refString="Horner JR, Goodwin MB. 2009. Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLOS ONE 4 (10): e 7626 DOI 10.1371 / journal. pone. 0007626." title="Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus" type="journal article" year="2009">
|
||
<emphasis box="[1058,1356,951,978]" italics="true" pageId="43" pageNumber="44">Horner & Goodwin, 2009</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,1017,1645]" box="[498,774,1017,1045]" pageId="43" pageNumber="44">
|
||
<heading bold="true" box="[498,774,1017,1045]" fontSize="11" level="3" pageId="43" pageNumber="44" reason="6">
|
||
<emphasis bold="true" box="[498,774,1017,1045]" italics="true" pageId="43" pageNumber="44">Palate and quadrate</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,1017,1645]" pageId="43" pageNumber="44">
|
||
The rostral process of the vomer is large in the less mature
|
||
<materialsCitation box="[1181,1413,1060,1087]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
, whereas it is small in the more mature
|
||
<materialsCitation box="[835,997,1100,1126]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
. The pits seen in the caudal process of
|
||
<materialsCitation collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
are regarded here as damage or individual variation, since they are not seen in any other
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[620,688,1181,1206]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="43" pageNumber="44" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[620,688,1181,1206]" italics="true" pageId="43" pageNumber="44">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens (
|
||
<materialsCitation box="[834,1071,1180,1207]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1084,1269,1180,1206]" collectionCode="BMRP" pageId="43" pageNumber="44" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1282,1400,1180,1206]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="UWBM" pageId="43" pageNumber="44" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
). The foramen in the ventral surface of the palatine in
|
||
<materialsCitation box="[1223,1457,1220,1247]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
is regarded here as either individual variation or a lesion, since this feature is not seen in any other specimen of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[719,788,1300,1325]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="43" pageNumber="44" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[719,788,1300,1325]" italics="true" pageId="43" pageNumber="44">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. The difference in texture of the caudal process (=anteroventral part of the pterygoid limb (
|
||
<bibRefCitation author="Molnar RE" box="[823,981,1339,1366]" journalOrPublisher="Palaeontographica Abteilung A" pageId="43" pageNumber="44" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[823,981,1339,1366]" italics="true" pageId="43" pageNumber="44">Molnar, 1990</emphasis>
|
||
</bibRefCitation>
|
||
)) of the ectopterygoid is not congruent with the growth series found here, which follows a sequence from
|
||
<materialsCitation box="[1214,1366,1379,1405]" collectionCode="SDSM" pageId="43" pageNumber="44" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
(not rough) to
|
||
<materialsCitation box="[498,658,1419,1445]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
(rough) to
|
||
<materialsCitation box="[797,913,1419,1445]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
(not rough). The thickness of the quadrate process of the pterygoid is congruent with the growth series, which is thick in the subadult category (
|
||
<materialsCitation box="[611,775,1499,1526]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
), whereas it is thin in adults (e.g.,
|
||
<materialsCitation box="[1168,1328,1499,1525]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1339,1456,1499,1525]" collectionCode="MOR" pageId="43" pageNumber="44" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="SDSM" pageId="43" pageNumber="44" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
). A possible ontogenetic progression is seen on the medial surface of the orbital process of the quadrate, which is coarse in the less mature
|
||
<materialsCitation box="[1195,1350,1579,1605]" collectionCode="SDSM" pageId="43" pageNumber="44" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
, whereas it is smooth in the more mature
|
||
<materialsCitation box="[834,997,1618,1644]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,1683,1913]" box="[498,636,1683,1711]" pageId="43" pageNumber="44">
|
||
<heading bold="true" box="[498,636,1683,1711]" fontSize="11" level="3" pageId="43" pageNumber="44" reason="6">
|
||
<emphasis bold="true" box="[498,636,1683,1711]" italics="true" pageId="43" pageNumber="44">Braincase</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="43.[498,1542,1683,1913]" lastBlockId="44.[498,1542,234,300]" lastPageId="44" lastPageNumber="45" pageId="43" pageNumber="44">
|
||
<bibRefCitation author="Molnar RE" box="[498,668,1727,1753]" journalOrPublisher="Palaeontographica Abteilung A" pageId="43" pageNumber="44" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[498,668,1727,1753]" italics="true" pageId="43" pageNumber="44">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
reported that the joint surface for the laterosphenoid on the postorbital is variable, where it is convex (
|
||
<materialsCitation box="[861,1101,1767,1794]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
), shallowly concave (TMP 1981.012.0001), or deeply concave (
|
||
<materialsCitation box="[918,1084,1807,1833]" collectionCode="LACM" pageId="43" pageNumber="44" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
). However, examination of
|
||
<materialsCitation collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="43" pageNumber="44" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
for this study found that the joint surface is concave. It is possible that the difference between shallow and deep states is ontogenetically informative, but the rim that surrounds the joint surface is often damaged and missing, preventing a precise assessment of depth.
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,337,1204]" box="[498,748,337,365]" pageId="44" pageNumber="45">
|
||
<heading bold="true" box="[498,748,337,365]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="6">
|
||
<emphasis bold="true" box="[498,748,337,365]" italics="true" pageId="44" pageNumber="45">Mandibular ramus</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,337,1204]" pageId="44" pageNumber="45">
|
||
Molnar suggested that the intercoronoid is fused to the dentary in most specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,565,421,446]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,565,421,446]" italics="true" pageId="44" pageNumber="45">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, but that condition has not been seen during the course of this study; in all cases, the bones are separate.
|
||
<bibRefCitation author="Molnar RE" box="[722,889,460,486]" journalOrPublisher="Palaeontographica Abteilung A" pageId="44" pageNumber="45" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[722,889,460,486]" italics="true" pageId="44" pageNumber="45">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
is correct in stating that the right angular of
|
||
<materialsCitation box="[1403,1517,460,486]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
is coarsened by lesions, and so this is not a growth-related change. The fusion of the prearticular reported by
|
||
<bibRefCitation author="Molnar RE" box="[788,957,540,566]" journalOrPublisher="Palaeontographica Abteilung A" pageId="44" pageNumber="45" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[788,957,540,566]" italics="true" pageId="44" pageNumber="45">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<materialsCitation box="[998,1114,540,566]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
might be a growth-related coossification event. The thickened surangular shelf seen in
|
||
<materialsCitation box="[1170,1330,580,606]" collectionCode="LACM" pageId="44" pageNumber="45" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
, in contrast to the thinner condition reported in
|
||
<materialsCitation box="[855,1088,619,646]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="44" pageNumber="45" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Molnar RE" box="[1107,1264,619,646]" journalOrPublisher="Palaeontographica Abteilung A" pageId="44" pageNumber="45" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[1107,1264,619,646]" italics="true" pageId="44" pageNumber="45">Molnar, 1990</emphasis>
|
||
</bibRefCitation>
|
||
), might be another growth-related change.
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,337,1204]" pageId="44" pageNumber="45">
|
||
<bibRefCitation author="Molnar RE" box="[530,697,699,725]" journalOrPublisher="Palaeontographica Abteilung A" pageId="44" pageNumber="45" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[530,697,699,725]" italics="true" pageId="44" pageNumber="45">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
noted the presence of perforations in the lateral plate of the surangular in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,567,740,765]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,740,765]" italics="true" pageId="44" pageNumber="45">T. rex</emphasis>
|
||
</taxonomicName>
|
||
; these purportedly pathological features (
|
||
<bibRefCitation author="Wolff EDS & Salisbury SW & Horner JR & Varricchio DJ" box="[1058,1258,739,765]" journalOrPublisher="PLOS ONE" pageId="44" pageNumber="45" pagination="e 7288" part="4" refId="ref56746" refString="Wolff EDS, Salisbury SW, Horner JR, Varricchio DJ. 2009. Common avian infection plagued the tyrant dinosaurs. PLOS ONE 4 (9): e 7288 DOI 10.1371 / journal. pone. 0007288." title="Common avian infection plagued the tyrant dinosaurs" type="journal article" year="2009">
|
||
<emphasis box="[1058,1258,739,765]" italics="true" pageId="44" pageNumber="45">Wolff et al., 2009</emphasis>
|
||
</bibRefCitation>
|
||
) deserve comment. Three perforations occur in the same location in different specimens. The first penetrates the bone rostroventral to the caudal surangular foramen (csf) at the dorsal margin of the angular; this is seen in two young adults (
|
||
<materialsCitation box="[1036,1171,858,885]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1184,1357,859,885]" collectionCode="UWBM" pageId="44" pageNumber="45" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
) and in one adult (
|
||
<materialsCitation box="[573,695,899,925]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
). The second penetrates the region ahead of the csf below the surangular shelf; this is seen in one young adult (
|
||
<materialsCitation box="[939,1115,938,964]" collectionCode="UWBM" pageId="44" pageNumber="45" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
), three adults (
|
||
<materialsCitation box="[1288,1408,938,964]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1420,1537,938,964]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,646,978,1005]" collectionCode="RSM" pageId="44" pageNumber="45" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
), and in the senescent adult (
|
||
<materialsCitation box="[991,1183,978,1004]" collectionCode="FMNH" pageId="44" pageNumber="45" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
). The third penetrates the rostroventral quadrant of the bone ahead of the external mandibular fenestra, which is seen in one young adult (
|
||
<materialsCitation box="[792,968,1058,1084]" collectionCode="UWBM" pageId="44" pageNumber="45" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
), an adult (
|
||
<materialsCitation box="[1098,1219,1058,1084]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
), and in the senescent adult (
|
||
<materialsCitation box="[506,698,1098,1124]" collectionCode="FMNH" pageId="44" pageNumber="45" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
). Given the spotty distribution of these openings, and, indeed, their lesion-like appearance, they were not included in the cladistic analysis of growth; regardless, the regularity of their positions indicates that they might not be lesions.
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,1240,1589]" box="[498,702,1240,1268]" pageId="44" pageNumber="45">
|
||
<heading bold="true" box="[498,702,1240,1268]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="6">
|
||
<emphasis bold="true" box="[498,702,1240,1268]" italics="true" pageId="44" pageNumber="45">Suture closure</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,1240,1589]" pageId="44" pageNumber="45">
|
||
Molnar documented numerous cranial (postorbitojugal, quadratoquadratojugal, squamosoquadrate) and mandibular (angulosurangular, prearticulosurangular) suture closures in the adult
|
||
<materialsCitation box="[746,864,1363,1389]" collectionCode="MOR" pageId="44" pageNumber="45" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
, which he regarded as ontogenetic in nature. Given the relatively immature maturity of the specimen, and the fact that closure of these sutures are not seen in other
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[749,817,1444,1469]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[749,817,1444,1469]" italics="true" pageId="44" pageNumber="45">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and they are regarded here as individual variation; in at least one case, a clear lesion is seen at the point of fusion (e.g., right angulosurangular joint). Also, it is not obvious that the right quadratoquadratojugal suture is closed in the specimen.
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,1625,1934]" box="[498,991,1625,1653]" pageId="44" pageNumber="45">
|
||
<heading bold="true" box="[498,991,1625,1653]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="6">
|
||
<emphasis bold="true" box="[498,991,1625,1653]" italics="true" pageId="44" pageNumber="45">Ontogenetic variation among adults</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="44.[498,1542,1625,1934]" lastBlockId="45.[498,1542,683,749]" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45">
|
||
<bibRefCitation author="Molnar RE" box="[498,665,1669,1695]" journalOrPublisher="Palaeontographica Abteilung A" pageId="44" pageNumber="45" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[498,665,1669,1695]" italics="true" pageId="44" pageNumber="45">Molnar (1990)</emphasis>
|
||
</bibRefCitation>
|
||
regarded the texture of the subcutaneous surface of facial bones, inflation of cranial sinuses, joint fusions, development of the cornual process of the postorbital, as reflecting relative maturity. Also, he proposed that the variable shape of the maxillary fenestra and the joint surface for the laterosphenoid of the postorbital represent individual variation. The results here broadly agree with these assessments. He was less certain of the significance of the form of the premaxillary process of the nasal, difference in height of the palatal process of the maxilla and the form of the rostral plate of the vomer (
|
||
<bibRefCitation author="Molnar RE" box="[508,663,683,709]" journalOrPublisher="Palaeontographica Abteilung A" pageId="45" pageNumber="46" pagination="137 - 176" part="217" refId="ref55158" refString="Molnar RE. 1990. The cranial morphology of Tyrannosaurus rex. Palaeontographica Abteilung A 217: 137 - 176." title="The cranial morphology of Tyrannosaurus rex" type="journal article" year="1990">
|
||
<emphasis box="[508,663,683,709]" italics="true" pageId="45" pageNumber="46">Molnar, 1990</emphasis>
|
||
</bibRefCitation>
|
||
). The present study does not resolve the latter set of features, but they will be taken up in a future iteration of this work.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFE9FF85FDEBFF70339BFEF5" httpUri="http://table.plazi.org/id/DF5C84E2FFE9FF85FDEBFF70339BFEF5" pageId="45" pageNumber="46" startId="45.[525,580,248,269]" targetBox="[516,1524,349,600]" targetIsTable="true" targetPageId="45">
|
||
<paragraph blockId="45.[525,1515,248,329]" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[525,1426,248,270]" pageId="45" pageNumber="46">
|
||
Table 10 Data used in the correlation test of bite force and maturity in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1235,1420,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1235,1420,249,270]" italics="true" pageId="45" pageNumber="46">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The data for BMRP 2002.4.1are from
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[799,1062,277,299]" journalOrPublisher="Biology Letters" pageId="45" pageNumber="46" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[799,1062,277,299]" italics="true" pageId="45" pageNumber="46">Bates & Falkingham (2012)</emphasis>
|
||
</bibRefCitation>
|
||
and the data for the others are from
|
||
<bibRefCitation author="Gignac PM & Erickson GM" journalOrPublisher="Scientific Reports" pageId="45" pageNumber="46" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis italics="true" pageId="45" pageNumber="46">Gignac & Erickson (2017)</emphasis>
|
||
</bibRefCitation>
|
||
. Data used in the correlation test are in boldface.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="45" pageNumber="46">
|
||
<table box="[516,1524,349,600]" gridcols="5" gridrows="7" pageId="45" pageNumber="46">
|
||
<tr box="[516,1524,349,371]" gridrow="0" pageId="45" pageNumber="46">
|
||
<th box="[516,709,349,371]" gridcol="0" gridrow="0" pageId="45" pageNumber="46">Specimen</th>
|
||
<th box="[750,880,349,371]" gridcol="1" gridrow="0" pageId="45" pageNumber="46">Growth rank</th>
|
||
<th box="[921,1128,349,371]" gridcol="2" gridrow="0" pageId="45" pageNumber="46">Revised growth rank</th>
|
||
<th box="[1170,1333,349,371]" gridcol="3" gridrow="0" pageId="45" pageNumber="46">Bite force</th>
|
||
<th box="[1374,1524,349,371]" gridcol="4" gridrow="0" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1524,349,371]" pageId="45" pageNumber="46">Bite force rank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,390,412]" gridrow="1" pageId="45" pageNumber="46">
|
||
<th box="[516,709,390,412]" gridcol="0" gridrow="1" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,665,390,412]" collectionCode="BMRP" pageId="45" pageNumber="46" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,390,412]" gridcol="1" gridrow="1" pageId="45" pageNumber="46">4</td>
|
||
<td box="[921,1128,390,412]" gridcol="2" gridrow="1" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,390,411]" pageId="45" pageNumber="46">1</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,390,412]" gridcol="3" gridrow="1" pageId="45" pageNumber="46">
|
||
2,400
|
||
<emphasis box="[1221,1233,390,412]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
3,850 N
|
||
</td>
|
||
<td box="[1374,1524,390,412]" gridcol="4" gridrow="1" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,390,411]" pageId="45" pageNumber="46">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,428,450]" gridrow="2" pageId="45" pageNumber="46">
|
||
<th box="[516,709,428,450]" gridcol="0" gridrow="2" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,709,428,450]" collectionCode="TMP" pageId="45" pageNumber="46" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,428,450]" gridcol="1" gridrow="2" pageId="45" pageNumber="46">8</td>
|
||
<td box="[921,1128,428,450]" gridcol="2" gridrow="2" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,428,449]" pageId="45" pageNumber="46">2</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,428,450]" gridcol="3" gridrow="2" pageId="45" pageNumber="46">
|
||
12,197
|
||
<emphasis box="[1233,1245,428,450]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
21,799 N
|
||
</td>
|
||
<td box="[1374,1524,428,450]" gridcol="4" gridrow="2" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,428,449]" pageId="45" pageNumber="46">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,465,487]" gridrow="3" pageId="45" pageNumber="46">
|
||
<th box="[516,709,465,487]" gridcol="0" gridrow="3" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,610,466,487]" collectionCode="MOR" pageId="45" pageNumber="46" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,465,487]" gridcol="1" gridrow="3" pageId="45" pageNumber="46">16</td>
|
||
<td box="[921,1128,465,487]" gridcol="2" gridrow="3" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,465,486]" pageId="45" pageNumber="46">3</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,465,487]" gridcol="3" gridrow="3" pageId="45" pageNumber="46">
|
||
14,201
|
||
<emphasis box="[1233,1245,465,487]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
30,487 N
|
||
</td>
|
||
<td box="[1374,1524,465,487]" gridcol="4" gridrow="3" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,465,486]" pageId="45" pageNumber="46">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,503,525]" gridrow="4" pageId="45" pageNumber="46">
|
||
<th box="[516,709,503,525]" gridcol="0" gridrow="4" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,646,503,524]" collectionCode="LACM" pageId="45" pageNumber="46" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,503,525]" gridcol="1" gridrow="4" pageId="45" pageNumber="46">17</td>
|
||
<td box="[921,1128,503,525]" gridcol="2" gridrow="4" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,503,524]" pageId="45" pageNumber="46">4</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,503,525]" gridcol="3" gridrow="4" pageId="45" pageNumber="46">
|
||
16,352
|
||
<emphasis box="[1233,1245,503,525]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
31,284 N
|
||
</td>
|
||
<td box="[1374,1524,503,525]" gridcol="4" gridrow="4" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,503,524]" pageId="45" pageNumber="46">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,540,563]" gridrow="5" pageId="45" pageNumber="46">
|
||
<th box="[516,709,540,563]" gridcol="0" gridrow="5" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,610,541,562]" collectionCode="MOR" pageId="45" pageNumber="46" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,540,563]" gridcol="1" gridrow="5" pageId="45" pageNumber="46">18</td>
|
||
<td box="[921,1128,540,563]" gridcol="2" gridrow="5" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,541,562]" pageId="45" pageNumber="46">5</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,540,563]" gridcol="3" gridrow="5" pageId="45" pageNumber="46">
|
||
13,736
|
||
<emphasis box="[1233,1245,541,563]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
28,101 N
|
||
</td>
|
||
<td box="[1374,1524,540,563]" gridcol="4" gridrow="5" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,540,561]" pageId="45" pageNumber="46">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,578,600]" gridrow="6" pageId="45" pageNumber="46">
|
||
<th box="[516,709,578,600]" gridcol="0" gridrow="6" pageId="45" pageNumber="46">
|
||
<materialsCitation box="[516,667,578,600]" collectionCode="FMNH" pageId="45" pageNumber="46" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[750,880,578,600]" gridcol="1" gridrow="6" pageId="45" pageNumber="46">20</td>
|
||
<td box="[921,1128,578,600]" gridcol="2" gridrow="6" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[921,933,578,599]" pageId="45" pageNumber="46">6</emphasis>
|
||
</td>
|
||
<td box="[1170,1333,578,600]" gridcol="3" gridrow="6" pageId="45" pageNumber="46">
|
||
17,769
|
||
<emphasis box="[1233,1245,578,600]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
34,522 N
|
||
</td>
|
||
<td box="[1374,1524,578,600]" gridcol="4" gridrow="6" pageId="45" pageNumber="46">
|
||
<emphasis bold="true" box="[1374,1386,578,599]" pageId="45" pageNumber="46">6</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="45.[498,1542,791,1421]" box="[498,905,791,821]" pageId="45" pageNumber="46">
|
||
<heading bold="true" box="[498,905,791,821]" fontSize="12" level="2" pageId="45" pageNumber="46" reason="0">
|
||
<emphasis bold="true" box="[498,905,791,821]" pageId="45" pageNumber="46">Secondary metamorphosis</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="45.[498,1542,791,1421]" pageId="45" pageNumber="46">
|
||
The abrupt, 2-year transition in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[881,950,838,863]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[881,950,838,863]" italics="true" pageId="45" pageNumber="46">T. rex</emphasis>
|
||
</taxonomicName>
|
||
from the sleek craniomandibular skeleton of juveniles to the deep and stout form of subadults (a change that is terminal) is an example of secondary metamorphosis, an extreme transformation of morphology that is associated with sexual maturity (
|
||
<bibRefCitation author="Rose CS & Reiss JO" box="[883,1100,957,983]" editor="Hanken H & Hall BK" journalOrPublisher="Chicago: University of Chicago Press" pageId="45" pageNumber="46" refId="ref55796" refString="Rose CS, Reiss JO. 1993. Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms. In: Hanken H, Hall BK, eds. The Skull: Development. Vol. 1. Chicago: University of Chicago Press." title="Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms" type="book" volumeTitle="The Skull: Development. Vol. 1" year="1993">
|
||
<emphasis box="[883,1100,957,983]" italics="true" pageId="45" pageNumber="46">Rose & Reiss, 1993</emphasis>
|
||
</bibRefCitation>
|
||
). Every bone and anatomical domain (e.g., paranasal air sac system, dentition, skull frame, musculature, integumentary system, etc.) is involved in reshaping the entire head skeleton, making juveniles and mature specimens so different that they have been taken to be different taxa (
|
||
<bibRefCitation author="Molnar RE" box="[1381,1536,1076,1103]" journalOrPublisher="Journal of Paleontology" pageId="45" pageNumber="46" pagination="102 - 108" part="54" refId="ref55134" refString="Molnar RE. 1980. An albertosaur from the Hell Creek Formation of Montana. Journal of Paleontology 54: 102 - 108." title="An albertosaur from the Hell Creek Formation of Montana" type="journal article" year="1980">
|
||
<emphasis box="[1381,1536,1076,1103]" italics="true" pageId="45" pageNumber="46">Molnar, 1980</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Bakker RT & Williams M & Currie PJ" box="[498,873,1116,1143]" journalOrPublisher="Hunteria" pageId="45" pageNumber="46" pagination="1 - 30" part="1" refId="ref52030" refString="Bakker RT, Williams M, Currie PJ. 1988. Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana. Hunteria 1: 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana" type="journal article" year="1988">
|
||
<emphasis box="[498,873,1116,1143]" italics="true" pageId="45" pageNumber="46">Bakker, Williams & Currie, 1988</emphasis>
|
||
</bibRefCitation>
|
||
). The occurrence of multiple transformative events across the head skeleton (i.e., not a single event) is the primary evidence for metamorphosis (
|
||
<bibRefCitation author="Rose CS & Reiss JO" box="[508,727,1196,1222]" editor="Hanken H & Hall BK" journalOrPublisher="Chicago: University of Chicago Press" pageId="45" pageNumber="46" refId="ref55796" refString="Rose CS, Reiss JO. 1993. Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms. In: Hanken H, Hall BK, eds. The Skull: Development. Vol. 1. Chicago: University of Chicago Press." title="Metamorphosis and the vertebrate skull: ontogenetic patterns and developmental mechanisms" type="book" volumeTitle="The Skull: Development. Vol. 1" year="1993">
|
||
<emphasis box="[508,727,1196,1222]" italics="true" pageId="45" pageNumber="46">Rose & Reiss, 1993</emphasis>
|
||
</bibRefCitation>
|
||
). This case of secondary metamorphosis coincides with the trophic shift from juvenile to subadult, and, almost certainly, with sexual maturity, whereas it precedes somatic maturity (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[823,1054,1276,1302]" journalOrPublisher="Nature" pageId="45" pageNumber="46" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[823,1054,1276,1302]" italics="true" pageId="45" pageNumber="46">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
). This hypothesis can be rejected if, upon discovery of relatively complete large juvenile and subadult specimens, a quantitative comparison of bite force between large juveniles and subadults does not show a discontinuous increase in the magnitude of bite force between the growth categories.
|
||
</paragraph>
|
||
<paragraph blockId="45.[498,1542,1464,1935]" box="[498,843,1464,1494]" pageId="45" pageNumber="46">
|
||
<heading bold="true" box="[498,843,1464,1494]" fontSize="12" level="2" pageId="45" pageNumber="46" reason="0">
|
||
<emphasis bold="true" box="[498,843,1464,1494]" pageId="45" pageNumber="46">Bite force and maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="45.[498,1542,1464,1935]" pageId="45" pageNumber="46">
|
||
Different methods have been used to estimate bite forces of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1209,1278,1510,1535]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1209,1278,1510,1535]" italics="true" pageId="45" pageNumber="46">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, including adults (e.g.,
|
||
<bibRefCitation author="Erickson GM & Van Kirk SD & Su J & Levenston ME & Clear WE & Carter DR" box="[561,795,1549,1575]" journalOrPublisher="Nature" pageId="45" pageNumber="46" pagination="706 - 708" part="382" refId="ref53120" refString="Erickson GM, Van Kirk SD, Su J, Levenston ME, Clear WE, Carter DR. 1996. Bite-force estimation for Tyrannosaurus rex from tooth-marked bones. Nature 382 (6593): 706 - 708 DOI 10.1038 / 382706 a 0." title="Bite-force estimation for Tyrannosaurus rex from tooth-marked bones" type="journal article" year="1996">
|
||
<emphasis box="[561,795,1549,1575]" italics="true" pageId="45" pageNumber="46">Erickson et al., 1996</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[809,1097,1549,1576]" journalOrPublisher="Scientific Reports" pageId="45" pageNumber="46" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis box="[809,1097,1549,1576]" italics="true" pageId="45" pageNumber="46">Gignac & Erickson, 2017</emphasis>
|
||
</bibRefCitation>
|
||
) and a juvenile (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" journalOrPublisher="Biology Letters" pageId="45" pageNumber="46" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis italics="true" pageId="45" pageNumber="46">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<tableCitation box="[568,670,1589,1615]" captionStart="Table 10" captionStartId="45.[525,580,248,269]" captionTargetBox="[516,1524,349,600]" captionText="Table 10 Data used in the correlation test of bite force and maturity in Tyrannosaurus rex. The data for BMRP 2002.4.1are from Bates & Falkingham (2012) and the data for the others are from Gignac & Erickson (2017). Data used in the correlation test are in boldface." httpUri="http://table.plazi.org/id/DF5C84E2FFE9FF85FDEBFF70339BFEF5" pageId="45" pageNumber="46" tableUuid="DF5C84E2FFE9FF85FDEBFF70339BFEF5">Table 10</tableCitation>
|
||
). In each case, the maximum estimated bite force of the juvenile is consistently an order of magnitude lower than those for adults (
|
||
<tableCitation box="[1256,1358,1629,1655]" captionStart="Table 10" captionStartId="45.[525,580,248,269]" captionTargetBox="[516,1524,349,600]" captionText="Table 10 Data used in the correlation test of bite force and maturity in Tyrannosaurus rex. The data for BMRP 2002.4.1are from Bates & Falkingham (2012) and the data for the others are from Gignac & Erickson (2017). Data used in the correlation test are in boldface." httpUri="http://table.plazi.org/id/DF5C84E2FFE9FF85FDEBFF70339BFEF5" pageId="45" pageNumber="46" tableUuid="DF5C84E2FFE9FF85FDEBFF70339BFEF5">Table 10</tableCitation>
|
||
). The only exception to this is the allometric-scaling based estimate of
|
||
<bibRefCitation author="Meers MB" box="[1198,1349,1669,1695]" journalOrPublisher="Historical Biology" pageId="45" pageNumber="46" pagination="1 - 12" part="16" refId="ref55095" refString="Meers MB. 2002. Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior. Historical Biology 16 (1): 1 - 12 DOI 10.1080 / 0891296021000050755." title="Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior" type="journal article" year="2002">
|
||
<emphasis box="[1198,1349,1669,1695]" italics="true" pageId="45" pageNumber="46">Meers (2002)</emphasis>
|
||
</bibRefCitation>
|
||
for adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1467,1535,1670,1695]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1467,1535,1670,1695]" italics="true" pageId="45" pageNumber="46">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, which is an order of magnitude lower than the other estimates of maximum bite force that were obtained by other methods; regardless, the minimum estimate (7,600 N;
|
||
<bibRefCitation author="Meers MB" journalOrPublisher="Historical Biology" pageId="45" pageNumber="46" pagination="1 - 12" part="16" refId="ref55095" refString="Meers MB. 2002. Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior. Historical Biology 16 (1): 1 - 12 DOI 10.1080 / 0891296021000050755." title="Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior" type="journal article" year="2002">
|
||
<emphasis italics="true" pageId="45" pageNumber="46">Meers, 2002</emphasis>
|
||
</bibRefCitation>
|
||
) is greater than that of the maximum bite force of the juvenile (3,850 N;
|
||
<bibRefCitation author="Bates KT & Falkingham PL" journalOrPublisher="Biology Letters" pageId="45" pageNumber="46" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis italics="true" pageId="45" pageNumber="46">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="45.[498,1542,1464,1935]" lastBlockId="46.[498,1542,989,1892]" lastPageId="46" lastPageNumber="47" pageId="45" pageNumber="46">
|
||
Like crocodylians (
|
||
<emphasis box="[750,1024,1868,1895]" italics="true" pageId="45" pageNumber="46">Gignac & O’ Brien, 2016</emphasis>
|
||
),
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1047,1114,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1047,1114,1869,1894]" italics="true" pageId="45" pageNumber="46">T. rex</emphasis>
|
||
</taxonomicName>
|
||
has a high bite force for its body size (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[508,810,1908,1934]" journalOrPublisher="Biology Letters" pageId="45" pageNumber="46" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[508,810,1908,1934]" italics="true" pageId="45" pageNumber="46">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
). Bite force estimates for juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1201,1268,1909,1934]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1201,1268,1909,1934]" italics="true" pageId="45" pageNumber="46">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(2,600
|
||
<emphasis box="[1346,1361,1909,1934]" italics="true" pageId="45" pageNumber="46">–</emphasis>
|
||
4,010 N; BMRP 2002.4.1) are lower than those of adults (35,000
|
||
<emphasis box="[1060,1075,990,1015]" italics="true" pageId="46" pageNumber="47">–</emphasis>
|
||
57,000 N) and show a growth trend of positive allometry, when identical methods of bite force estimation have been used for both growth categories (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[786,1095,1069,1095]" journalOrPublisher="Biology Letters" pageId="46" pageNumber="47" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[786,1095,1069,1095]" italics="true" pageId="46" pageNumber="47">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
); that is, bite forces in adults are relatively and absolutely higher than those seen in juveniles.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5824998" ID-Zenodo-Dep="5824998" httpUri="https://zenodo.org/record/5824998/files/figure.png" pageId="46" pageNumber="47" startId="46.[524,589,736,758]" targetBox="[504,1452,238,702]" targetPageId="46">
|
||
<paragraph blockId="46.[524,1515,736,934]" pageId="46" pageNumber="47">
|
||
<emphasis bold="true" pageId="46" pageNumber="47">
|
||
Figure 13 Scatterplot showing the congruence in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1034,1222,737,758]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1034,1222,737,758]" italics="true" pageId="46" pageNumber="47">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
between bite force (i.e., and maturity).
|
||
</emphasis>
|
||
Growth stage rank, corresponding to the increasing sequence of nodes in Fig. 2, is along the
|
||
<emphasis box="[559,570,796,817]" italics="true" pageId="46" pageNumber="47">x</emphasis>
|
||
-axis; growth stage rank refers to the relative maturity of the specimens for which bite force has been estimated. Increasing bite force rank is along the
|
||
<emphasis box="[988,998,825,846]" italics="true" pageId="46" pageNumber="47">y</emphasis>
|
||
-axis; raw bite force data are from
|
||
<bibRefCitation author="Bates KT & Falkingham PL" journalOrPublisher="Biology Letters" pageId="46" pageNumber="47" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis italics="true" pageId="46" pageNumber="47">Bates & Falkingham (2012)</emphasis>
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[632,875,853,875]" journalOrPublisher="Scientific Reports" pageId="46" pageNumber="47" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis box="[632,875,853,875]" italics="true" pageId="46" pageNumber="47">Gignac & Erickson (2017)</emphasis>
|
||
</bibRefCitation>
|
||
. A Spearman correlation test on these data resulted in a significant correlation coefficient, indicating that bite force increases with maturity.
|
||
</paragraph>
|
||
<paragraph blockId="46.[524,1515,736,934]" box="[1099,1515,912,934]" pageId="46" pageNumber="47">Full-size DOI: 10.7717/peerj.9192/fig-13</paragraph>
|
||
</caption>
|
||
<paragraph blockId="46.[498,1542,989,1892]" pageId="46" pageNumber="47">
|
||
In light of these data, a comparison of bite force was made here between a sample of several adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[652,719,1189,1214]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[652,719,1189,1214]" italics="true" pageId="46" pageNumber="47">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[737,1022,1188,1215]" journalOrPublisher="Scientific Reports" pageId="46" pageNumber="47" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis box="[737,1022,1188,1215]" italics="true" pageId="46" pageNumber="47">Gignac & Erickson, 2017</emphasis>
|
||
</bibRefCitation>
|
||
) and a juvenile (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[1220,1526,1188,1215]" journalOrPublisher="Biology Letters" pageId="46" pageNumber="47" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[1220,1526,1188,1215]" italics="true" pageId="46" pageNumber="47">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
). Given that different methods were used to estimate bite force in the juvenile, on the one hand, and its adult counterparts on the other, the raw estimates were converted into ranks for a Spearman rank correlation test of the congruence between maturity and bite force (
|
||
<tableCitation box="[627,729,1348,1374]" captionStart="Table 10" captionStartId="45.[525,580,248,269]" captionTargetBox="[516,1524,349,600]" captionText="Table 10 Data used in the correlation test of bite force and maturity in Tyrannosaurus rex. The data for BMRP 2002.4.1are from Bates & Falkingham (2012) and the data for the others are from Gignac & Erickson (2017). Data used in the correlation test are in boldface." httpUri="http://table.plazi.org/id/DF5C84E2FFE9FF85FDEBFF70339BFEF5" pageId="46" pageNumber="47" tableUuid="DF5C84E2FFE9FF85FDEBFF70339BFEF5">Table 10</tableCitation>
|
||
). The data were tested for normality and a Shapiro-Wilk test; the growth ranks and bite force ranks are normally distributed (
|
||
<emphasis box="[1211,1225,1388,1413]" italics="true" pageId="46" pageNumber="47">p</emphasis>
|
||
= 0.961 for both). The Spearman rank correlation test resulted in a significant (
|
||
<emphasis box="[1219,1233,1428,1453]" italics="true" pageId="46" pageNumber="47">p</emphasis>
|
||
= 0.042) correlation coefficient (
|
||
<emphasis box="[634,645,1468,1493]" italics="true" pageId="46" pageNumber="47">
|
||
<emphasis box="[645,655,1480,1496]" italics="true" pageId="46" pageNumber="47">
|
||
<subScript attach="left" box="[645,655,1480,1496]" fontSize="7" pageId="46" pageNumber="47">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.829). Therefore, an ontogenetic progression of increasing bite force from juvenile to adult, and throughout adulthood (
|
||
<figureCitation box="[1033,1112,1507,1533]" captionStart="Figure 13" captionStartId="46.[524,589,736,758]" captionTargetBox="[504,1452,238,702]" captionTargetId="figure-367@46.[498,1459,234,709]" captionTargetPageId="46" captionText="Figure 13 Scatterplot showing the congruence in Tyrannosaurus rex between bite force (i.e., and maturity). Growth stage rank, corresponding to the increasing sequence of nodes in Fig. 2, is along the x-axis; growth stage rank refers to the relative maturity of the specimens for which bite force has been estimated. Increasing bite force rank is along the y-axis; raw bite force data are from Bates & Falkingham (2012) and Gignac & Erickson (2017). A Spearman correlation test on these data resulted in a significant correlation coefficient, indicating that bite force increases with maturity. Full-size DOI: 10.7717/peerj.9192/fig-13" figureDoi="http://doi.org/10.5281/zenodo.5824998" httpUri="https://zenodo.org/record/5824998/files/figure.png" pageId="46" pageNumber="47">Fig. 13</figureCitation>
|
||
), is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1253,1322,1508,1533]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1253,1322,1508,1533]" italics="true" pageId="46" pageNumber="47">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, which reflects the condition that is reported in crocodylians (
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[1007,1296,1547,1573]" journalOrPublisher="Journal of Zoology" pageId="46" pageNumber="47" pagination="132 - 142" part="295" refId="ref53694" refString="Gignac PM, Erickson GM. 2014. Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators. Journal of Zoology 295 (2): 132 - 142 DOI 10.1111 / jzo. 12187." title="Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators" type="journal article" year="2014">
|
||
<emphasis box="[1007,1296,1547,1573]" italics="true" pageId="46" pageNumber="47">Gignac & Erickson, 2014</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="46.[498,1542,989,1892]" lastBlockId="47.[498,1542,234,778]" lastPageId="47" lastPageNumber="48" pageId="46" pageNumber="47">
|
||
Tooth cross sectional shape serves as a proxy for bite force, where the tooth width to length ratio in crocodylians continuously increases with the ontogenetic increase in bite force (
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[622,903,1666,1693]" journalOrPublisher="Journal of Zoology" pageId="46" pageNumber="47" pagination="132 - 142" part="295" refId="ref53694" refString="Gignac PM, Erickson GM. 2014. Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators. Journal of Zoology 295 (2): 132 - 142 DOI 10.1111 / jzo. 12187." title="Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators" type="journal article" year="2014">
|
||
<emphasis box="[622,903,1666,1693]" italics="true" pageId="46" pageNumber="47">Gignac & Erickson, 2014</emphasis>
|
||
</bibRefCitation>
|
||
). In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[957,1023,1667,1692]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[957,1023,1667,1692]" italics="true" pageId="46" pageNumber="47">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, the tooth width to length ratio doubles from juveniles (e.g.,
|
||
<materialsCitation box="[672,857,1706,1733]" collectionCode="BMRP" pageId="46" pageNumber="47" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
, maxillary tooth 4, width to length ratio: 54%) to adults (e.g.,
|
||
<materialsCitation box="[559,675,1746,1772]" collectionCode="MOR" pageId="46" pageNumber="47" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
, maxillary tooth 4, width to length ratio: 98%), which might be evidence for a delay in the necessity for indenting and fracturing the bones of prey in juveniles, in contrast to the continuous pattern that is seen in crocodylians (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[1232,1536,1826,1852]" journalOrPublisher="Biology Letters" pageId="46" pageNumber="47" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[1232,1536,1826,1852]" italics="true" pageId="46" pageNumber="47">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[498,787,1866,1892]" journalOrPublisher="Journal of Zoology" pageId="46" pageNumber="47" pagination="132 - 142" part="295" refId="ref53694" refString="Gignac PM, Erickson GM. 2014. Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators. Journal of Zoology 295 (2): 132 - 142 DOI 10.1111 / jzo. 12187." title="Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators" type="journal article" year="2014">
|
||
<emphasis box="[498,787,1866,1892]" italics="true" pageId="46" pageNumber="47">Gignac & Erickson, 2014</emphasis>
|
||
</bibRefCitation>
|
||
). Ergo, a complete sample of bite force estimates across the metamorphic transition in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[810,876,235,260]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[810,876,235,260]" italics="true" pageId="47" pageNumber="48">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is required to test this hypothesis, which is based on gross morphology.
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,234,778]" pageId="47" pageNumber="48">
|
||
The relationship between bite force and size is linear in extant crocodylians, despite the great changes in the size and type of prey species, a clade that, unlike
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1326,1394,355,380]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1326,1394,355,380]" italics="true" pageId="47" pageNumber="48">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, does not undergo an abrupt, wholesale ontogenetic transformation in skull shape (
|
||
<bibRefCitation author="Erickson GM & Lappin AK & Vliet KA" journalOrPublisher="Journal of Zoology" pageId="47" pageNumber="48" pagination="317 - 327" part="260" refId="ref53166" refString="Erickson GM, Lappin AK, Vliet KA. 2003. The ontogeny of bite-force performance in American Alligator (Alligator mississippiensis). Journal of Zoology 260 (3): 317 - 327 DOI 10.1017 / S 0952836903003819." title="The ontogeny of bite-force performance in American Alligator (Alligator mississippiensis)" type="journal article" year="2003">
|
||
<emphasis italics="true" pageId="47" pageNumber="48">Erickson, Lappin & Vliet, 2003</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Erickson GM & Gignac PM & Lappin AK & Vliet KA & Brueggen JD & Webb GJW" box="[759,994,433,460]" journalOrPublisher="Journal of Zoology" pageId="47" pageNumber="48" pagination="48 - 55" part="292" refId="ref53338" refString="Erickson GM, Gignac PM, Lappin AK, Vliet KA, Brueggen JD, Webb GJW. 2013. A comparative analysis of ontogenetic bite-force scaling among Crocodylia. Journal of Zoology 292 (1): 48 - 55 DOI 10.1111 / jzo. 12081." title="A comparative analysis of ontogenetic bite-force scaling among Crocodylia" type="journal article" year="2013">
|
||
<emphasis box="[759,994,433,460]" italics="true" pageId="47" pageNumber="48">Erickson et al., 2013</emphasis>
|
||
</bibRefCitation>
|
||
). If bite forces in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1207,1276,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1207,1276,434,459]" italics="true" pageId="47" pageNumber="48">T. rex</emphasis>
|
||
</taxonomicName>
|
||
do increase abruptly across metamorphosis, then this is fundamentally unlike the situation in crocodylians, where bite force matches or exceeds prey shear forces in crocodylians (
|
||
<bibRefCitation author="Gignac PM & Erickson GM" journalOrPublisher="Journal of Zoology" pageId="47" pageNumber="48" pagination="132 - 142" part="295" refId="ref53694" refString="Gignac PM, Erickson GM. 2014. Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators. Journal of Zoology 295 (2): 132 - 142 DOI 10.1111 / jzo. 12187." title="Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators" type="journal article" year="2014">
|
||
<emphasis italics="true" pageId="47" pageNumber="48">Gignac & Erickson, 2014</emphasis>
|
||
</bibRefCitation>
|
||
). In contrast, apical tooth pressures in the dinosaur will be found to not equal or exceed the maximum shear strength of significantly larger prey until metamorphosis has occurred. If true, then the ontogenetic change reflects a discontinuous and abrupt dietary change to significantly larger prey that was necessary to sustain the metabolic demands of a 1.8-tonne animal on its way to becoming a 3+ tonne predator.
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,812,1480]" box="[498,751,812,840]" pageId="47" pageNumber="48">
|
||
<heading bold="true" box="[498,751,812,840]" fontSize="11" level="3" pageId="47" pageNumber="48" reason="6">
|
||
<emphasis bold="true" box="[498,751,812,840]" italics="true" pageId="47" pageNumber="48">Tooth morphology</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,812,1480]" pageId="47" pageNumber="48">
|
||
A comparison of tooth width to length ratios gives some indication of whether or not an abrupt, discontinuous increase in bite force is reasonable to expect in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1324,1392,896,921]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1324,1392,896,921]" italics="true" pageId="47" pageNumber="48">T. rex</emphasis>
|
||
</taxonomicName>
|
||
growth. The comparisons made here are based on the lateral teeth of the maxilla and dentary; that is, excluding the incisiform first two teeth of the maxilla and the two mesial subconical teeth of the dentary that have higher width to length ratios than the successive ziphiform teeth. In large juveniles (
|
||
<materialsCitation box="[902,1087,1055,1081]" collectionCode="BMRP" pageId="47" pageNumber="48" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
,
|
||
<emphasis box="[1099,1115,1056,1081]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 34) the mean width to length ratio across all teeth is 54%; in young adults (
|
||
<materialsCitation box="[973,1107,1095,1122]" collectionCode="MOR" pageId="47" pageNumber="48" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
,
|
||
<emphasis box="[1120,1136,1096,1121]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 20) the ratio is 68%, and in adults (
|
||
<materialsCitation box="[584,703,1135,1161]" collectionCode="MOR" pageId="47" pageNumber="48" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<emphasis box="[715,731,1135,1160]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 14;
|
||
<materialsCitation box="[804,965,1135,1161]" collectionCode="LACM" pageId="47" pageNumber="48" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<emphasis box="[977,993,1135,1160]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 6;
|
||
<materialsCitation box="[1052,1169,1135,1161]" collectionCode="MOR" pageId="47" pageNumber="48" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<emphasis box="[1181,1197,1135,1160]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 2;
|
||
<materialsCitation box="[1257,1401,1134,1161]" collectionCode="RSM" pageId="47" pageNumber="48" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
,
|
||
<emphasis box="[1412,1428,1135,1160]" italics="true" pageId="47" pageNumber="48">n</emphasis>
|
||
= 10) the ratio is 79%, 71%, 85%, and 85%, respectively (
|
||
<tableCitation box="[1056,1157,1174,1201]" captionStart="Table 11" captionStartId="48.[117,172,248,269]" captionTargetBox="[108,1525,349,1723]" captionText="Table 11 Summary of width to length ratios of maxillary and dentary teeth in adult Tyrannosaurus rex compared with representative adults of other tyrannosaurids. In T. rex the maxillary teeth tend to be wider than in other tyrannosaurids, and also at the mesial end of the tooth row of the dentary. However, the characterization by Osborn (1906) of the teeth in T. rex as generally wider than long is not supported by these data." httpUri="http://table.plazi.org/id/DF5C84E2FFF4FF98FF93FF7032AAFEF5" pageId="47" pageNumber="48" tableUuid="DF5C84E2FFF4FF98FF93FF7032AAFEF5">Table 11</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,812,1480]" pageId="47" pageNumber="48">
|
||
Bite force estimates for the juvenile are 2,400 N for the mesial teeth and 3,850 N for the distal teeth (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[691,1000,1254,1281]" journalOrPublisher="Biology Letters" pageId="47" pageNumber="48" pagination="20180160" part="14" refId="ref52146" refString="Bates KT, Falkingham PL. 2018. Correction to ' estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics '. Biology Letters 14 (4): 20180160 DOI 10.1098 / rsbl. 2018.0160." title="Correction to ' estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2018">
|
||
<emphasis box="[691,1000,1254,1281]" italics="true" pageId="47" pageNumber="48">Bates & Falkingham, 2018</emphasis>
|
||
</bibRefCitation>
|
||
); the minimum bite force estimates for a subadult (TMP 1981.006.0001) is 12,197 N, and in adults (
|
||
<materialsCitation box="[1191,1312,1294,1320]" collectionCode="MOR" pageId="47" pageNumber="48" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
, LACM 23944,
|
||
<materialsCitation box="[498,617,1334,1360]" collectionCode="MOR" pageId="47" pageNumber="48" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
) the estimates are 13,736 N, 16,352 N, and 14,201 N, respectively (
|
||
<bibRefCitation author="Gignac PM & Erickson GM" journalOrPublisher="Scientific Reports" pageId="47" pageNumber="48" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis italics="true" pageId="47" pageNumber="48">Gignac & Erickson, 2017</emphasis>
|
||
</bibRefCitation>
|
||
). Therefore, the order-of-magnitude increase in bite force, and the 12% increase in tooth width that is seen between juveniles and subadults, are naively predicted here to occur abruptly in the subadult stage, no later than 15 years old.
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,1519,1751]" box="[498,878,1519,1549]" pageId="47" pageNumber="48">
|
||
<heading bold="true" box="[498,878,1519,1549]" fontSize="12" level="2" pageId="47" pageNumber="48" reason="0">
|
||
<emphasis bold="true" box="[498,878,1519,1549]" pageId="47" pageNumber="48">Tooth count and maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,1519,1751]" pageId="47" pageNumber="48">
|
||
In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[530,596,1566,1591]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="47" pageNumber="48" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[530,596,1566,1591]" italics="true" pageId="47" pageNumber="48">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, tooth count in the maxilla and dentary initially increases, and then decreases over the course of the growth series. Despite this general trend, variation is seen from the young adult growth stage onwards; therefore, the congruence between maturity and the tooth number in the maxilla and the dentary was tested using Spearman rank correlation, assuming a null hypothesis of noncorrelation.
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,1784,1894]" box="[498,1019,1784,1812]" pageId="47" pageNumber="48">
|
||
<heading bold="true" box="[498,1019,1784,1812]" fontSize="11" level="3" pageId="47" pageNumber="48" reason="6">
|
||
<emphasis bold="true" box="[498,1019,1784,1812]" italics="true" pageId="47" pageNumber="48">Correlation with maxillary tooth count</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="47.[498,1542,1784,1894]" lastBlockId="48.[498,1542,1780,1926]" lastPageId="48" lastPageNumber="49" pageId="47" pageNumber="48">
|
||
The bivariate scatterplot of ranked data (
|
||
<figureCitation box="[982,1062,1828,1854]" captionStart="Figure 14" captionStartId="49.[524,589,706,728]" captionTargetBox="[506,1454,236,674]" captionTargetId="figure-77@49.[498,1458,234,678]" captionTargetPageId="49" captionText="Figure 14 Bivariate scatterplot showing the relationship between maxillary tooth count with maturity among 14 specimens of Tyrannosaurus rex. Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which maxillary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Maxillary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-14" figureDoi="http://doi.org/10.5281/zenodo.5825000" httpUri="https://zenodo.org/record/5825000/files/figure.png" pageId="47" pageNumber="48">Fig. 14</figureCitation>
|
||
;
|
||
<tableCitation box="[1077,1178,1828,1854]" captionStart="Table 12" captionStartId="49.[525,580,958,979]" captionTargetBox="[516,1508,1058,1668]" captionText="Table 12 Summary of maxillary tooth count data and growth rank in Tyrannosaurus rex. Summary of raw and ranked data used for the Spearman correlation test between maturity and maxillary tooth count in T. rex. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FFF5FF99FDEBFC3633DDFBB2" pageId="47" pageNumber="48" tableUuid="DF5C84E2FFF5FF99FDEBFC3633DDFBB2">Table 12</tableCitation>
|
||
) shows that maxillary tooth number initially increases between the first two growth stage ranks (from 15 to 16 teeth) before it abruptly decreases (from 16 to 12 teeth) at the third rank (
|
||
<figureCitation box="[1292,1371,1780,1806]" captionStart="Figure 14" captionStartId="49.[524,589,706,728]" captionTargetBox="[506,1454,236,674]" captionTargetId="figure-77@49.[498,1458,234,678]" captionTargetPageId="49" captionText="Figure 14 Bivariate scatterplot showing the relationship between maxillary tooth count with maturity among 14 specimens of Tyrannosaurus rex. Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which maxillary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Maxillary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-14" figureDoi="http://doi.org/10.5281/zenodo.5825000" httpUri="https://zenodo.org/record/5825000/files/figure.png" pageId="48" pageNumber="49">Fig. 14</figureCitation>
|
||
). Tooth count rank is constant until the sixth rank, where variation, toward a lower tooth count (from 12 to 11 teeth), is first seen (
|
||
<figureCitation box="[905,983,1859,1885]" captionStart="Figure 14" captionStartId="49.[524,589,706,728]" captionTargetBox="[506,1454,236,674]" captionTargetId="figure-77@49.[498,1458,234,678]" captionTargetPageId="49" captionText="Figure 14 Bivariate scatterplot showing the relationship between maxillary tooth count with maturity among 14 specimens of Tyrannosaurus rex. Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which maxillary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Maxillary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-14" figureDoi="http://doi.org/10.5281/zenodo.5825000" httpUri="https://zenodo.org/record/5825000/files/figure.png" pageId="48" pageNumber="49">Fig. 14</figureCitation>
|
||
). Prima facie, tooth loss covaries with maturity, as is seen in other tyrannosaurids (
|
||
<bibRefCitation author="Carr TD & Varricchio DJ & Sedlmayr JC & Roberts EM & Moore JR" box="[913,1103,1899,1926]" journalOrPublisher="Scientific Reports" pageId="48" pageNumber="49" pagination="44942" part="7" refId="ref52780" refString="Carr TD, Varricchio DJ, Sedlmayr JC, Roberts EM, Moore JR. 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7 (1): 44942 DOI 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">
|
||
<emphasis box="[913,1103,1899,1926]" italics="true" pageId="48" pageNumber="49">Carr et al., 2017</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFF4FF98FF93FF7032AAFEF5" httpUri="http://table.plazi.org/id/DF5C84E2FFF4FF98FF93FF7032AAFEF5" pageId="48" pageNumber="49" startId="48.[117,172,248,269]" targetBox="[108,1525,349,1723]" targetIsTable="true" targetPageId="48">
|
||
<paragraph blockId="48.[117,1515,248,329]" pageId="48" pageNumber="49">
|
||
<emphasis bold="true" pageId="48" pageNumber="49">
|
||
Table 11 Summary of width to length ratios of maxillary and dentary teeth in adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[942,1126,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[942,1126,249,270]" italics="true" pageId="48" pageNumber="49">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
compared with representative adults of other tyrannosaurids.
|
||
</emphasis>
|
||
In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[364,418,278,299]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[364,418,278,299]" italics="true" pageId="48" pageNumber="49">T. rex</emphasis>
|
||
</taxonomicName>
|
||
the maxillary teeth tend to be wider than in other tyrannosaurids, and also at the mesial end of the tooth row of the dentary. However, the characterization by
|
||
<bibRefCitation author="Osborn HF" box="[528,665,306,328]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="48" pageNumber="49" pagination="281 - 296" part="22" refId="ref55443" refString="Osborn HF. 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication). Bulletin of the American Museum of Natural History 22: 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication)" type="journal article" year="1906">
|
||
<emphasis box="[528,665,306,328]" italics="true" pageId="48" pageNumber="49">Osborn (1906)</emphasis>
|
||
</bibRefCitation>
|
||
of the teeth in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[815,871,307,328]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[815,871,307,328]" italics="true" pageId="48" pageNumber="49">T. rex</emphasis>
|
||
</taxonomicName>
|
||
as generally wider than long is not supported by these data.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="48" pageNumber="49">
|
||
<table box="[108,1525,349,1723]" gridcols="11" gridrows="35" pageId="48" pageNumber="49">
|
||
<tr box="[108,1525,349,371]" gridrow="0" pageId="48" pageNumber="49" rowspan-0="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1">
|
||
<th box="[212,272,349,371]" gridcol="1" gridrow="0" pageId="48" pageNumber="49">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[212,272,349,370]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
|
||
</th>
|
||
<th box="[1073,1172,349,371]" gridcol="8" gridrow="0" pageId="48" pageNumber="49">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1073,1172,349,370]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="torosus">D. torosus</taxonomicName>
|
||
</th>
|
||
<th box="[1224,1327,349,371]" gridcol="9" gridrow="0" pageId="48" pageNumber="49">
|
||
<taxonomicName box="[1224,1327,349,371]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="horneri">D. horneri</taxonomicName>
|
||
</th>
|
||
<taxonomicName authority="MOR" authorityName="MOR" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<th box="[1376,1525,349,371]" gridcol="10" gridrow="0" pageId="48" pageNumber="49">
|
||
<emphasis bold="true" box="[1376,1525,349,371]" italics="true" pageId="48" pageNumber="49">A. sarcophagus</emphasis>
|
||
</th>
|
||
<tr box="[108,1525,398,449]" gridrow="1" pageId="48" pageNumber="49" rowspan-0="1">
|
||
<td box="[212,272,398,449]" gridcol="1" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[212,267,398,448]" collectionCode="MOR" pageId="48" pageNumber="49" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</td>
|
||
<td box="[326,392,398,449]" gridcol="2" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[326,392,398,448]" collectionCode="RSM" pageId="48" pageNumber="49" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</td>
|
||
<td box="[455,522,398,449]" gridcol="3" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[455,522,398,448]" collectionCode="LACM" pageId="48" pageNumber="49" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</td>
|
||
<td box="[584,639,398,449]" gridcol="4" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[584,639,398,448]" collectionCode="MOR" pageId="48" pageNumber="49" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</td>
|
||
<td box="[698,750,398,449]" gridcol="5" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[698,746,398,448]" collectionCode="CM" pageId="48" pageNumber="49" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</td>
|
||
<td box="[811,870,398,449]" gridcol="6" gridrow="1" pageId="48" pageNumber="49">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[811,870,398,419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="48" pageNumber="49" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[811,870,398,419]" italics="true" pageId="48" pageNumber="49">T. rex</emphasis>
|
||
</taxonomicName>
|
||
mean
|
||
</td>
|
||
<td box="[924,1019,398,449]" gridcol="7" gridrow="1" pageId="48" pageNumber="49">
|
||
<materialsCitation box="[924,1008,398,448]" collectionCode="BMRP" pageId="48" pageNumber="49" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</td>
|
||
<td box="[1073,1172,398,449]" gridcol="8" gridrow="1" pageId="48" pageNumber="49">CMN 8506</td>
|
||
<td box="[1224,1327,398,449]" gridcol="9" gridrow="1" pageId="48" pageNumber="49">MOR 1130</td>
|
||
<td box="[1376,1525,398,449]" gridcol="10" gridrow="1" pageId="48" pageNumber="49">
|
||
<emphasis bold="true" box="[1376,1427,398,419]" pageId="48" pageNumber="49">TMP</emphasis>
|
||
<emphasis bold="true" box="[1376,1519,427,448]" pageId="48" pageNumber="49">1981.010.0001</emphasis>
|
||
</td>
|
||
</tr>
|
||
</taxonomicName>
|
||
</tr>
|
||
<tr box="[108,1525,468,491]" gridrow="2" pageId="48" pageNumber="49">
|
||
<th box="[108,163,468,491]" gridcol="0" gridrow="2" pageId="48" pageNumber="49">Mx1</th>
|
||
<td box="[212,272,468,491]" gridcol="1" gridrow="2" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,469,491]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,468,491]" gridcol="2" gridrow="2" pageId="48" pageNumber="49">85%</td>
|
||
<td box="[455,522,468,491]" gridcol="3" gridrow="2" pageId="48" pageNumber="49">71%</td>
|
||
<td box="[584,639,468,491]" gridcol="4" gridrow="2" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,468,491]" gridcol="5" gridrow="2" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,469,491]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,468,491]" gridcol="6" gridrow="2" pageId="48" pageNumber="49">78%</td>
|
||
<td box="[924,1019,468,491]" gridcol="7" gridrow="2" pageId="48" pageNumber="49">?, 150%</td>
|
||
<td box="[1073,1172,468,491]" gridcol="8" gridrow="2" pageId="48" pageNumber="49">87%</td>
|
||
<td box="[1224,1327,468,491]" gridcol="9" gridrow="2" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,468,491]" gridcol="10" gridrow="2" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,469,491]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,506,528]" gridrow="3" pageId="48" pageNumber="49">
|
||
<th box="[108,163,506,528]" gridcol="0" gridrow="3" pageId="48" pageNumber="49">Mx2</th>
|
||
<td box="[212,272,506,528]" gridcol="1" gridrow="3" pageId="48" pageNumber="49">–</td>
|
||
<td box="[326,392,506,528]" gridcol="2" gridrow="3" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,506,528]" gridcol="3" gridrow="3" pageId="48" pageNumber="49">
|
||
<emphasis box="[455,467,506,528]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[584,639,506,528]" gridcol="4" gridrow="3" pageId="48" pageNumber="49">91%</td>
|
||
<td box="[698,750,506,528]" gridcol="5" gridrow="3" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,506,528]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,506,528]" gridcol="6" gridrow="3" pageId="48" pageNumber="49">91%</td>
|
||
<td box="[924,1019,506,528]" gridcol="7" gridrow="3" pageId="48" pageNumber="49">73%, 83%</td>
|
||
<td box="[1073,1172,506,528]" gridcol="8" gridrow="3" pageId="48" pageNumber="49">64%</td>
|
||
<td box="[1224,1327,506,528]" gridcol="9" gridrow="3" pageId="48" pageNumber="49">79%</td>
|
||
<td box="[1376,1525,506,528]" gridcol="10" gridrow="3" pageId="48" pageNumber="49">56%</td>
|
||
</tr>
|
||
<tr box="[108,1525,544,566]" gridrow="4" pageId="48" pageNumber="49">
|
||
<th box="[108,163,544,566]" gridcol="0" gridrow="4" pageId="48" pageNumber="49">Mx3</th>
|
||
<td box="[212,272,544,566]" gridcol="1" gridrow="4" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,544,566]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,544,566]" gridcol="2" gridrow="4" pageId="48" pageNumber="49">65%</td>
|
||
<td box="[455,522,544,566]" gridcol="3" gridrow="4" pageId="48" pageNumber="49">69%</td>
|
||
<td box="[584,639,544,566]" gridcol="4" gridrow="4" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,544,566]" gridcol="5" gridrow="4" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,544,566]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,544,566]" gridcol="6" gridrow="4" pageId="48" pageNumber="49">67%</td>
|
||
<td box="[924,1019,544,566]" gridcol="7" gridrow="4" pageId="48" pageNumber="49">48%,?</td>
|
||
<td box="[1073,1172,544,566]" gridcol="8" gridrow="4" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,544,566]" gridcol="9" gridrow="4" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,544,566]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,544,566]" gridcol="10" gridrow="4" pageId="48" pageNumber="49">68%</td>
|
||
</tr>
|
||
<tr box="[108,1525,581,603]" gridrow="5" pageId="48" pageNumber="49">
|
||
<th box="[108,163,581,603]" gridcol="0" gridrow="5" pageId="48" pageNumber="49">Mx4</th>
|
||
<td box="[212,272,581,603]" gridcol="1" gridrow="5" pageId="48" pageNumber="49">98%</td>
|
||
<td box="[326,392,581,603]" gridcol="2" gridrow="5" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,581,603]" gridcol="3" gridrow="5" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,581,603]" gridcol="4" gridrow="5" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,581,603]" gridcol="5" gridrow="5" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,581,603]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,581,603]" gridcol="6" gridrow="5" pageId="48" pageNumber="49">98%</td>
|
||
<td box="[924,1019,581,603]" gridcol="7" gridrow="5" pageId="48" pageNumber="49">? 54%</td>
|
||
<td box="[1073,1172,581,603]" gridcol="8" gridrow="5" pageId="48" pageNumber="49">100%</td>
|
||
<td box="[1224,1327,581,603]" gridcol="9" gridrow="5" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,581,603]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,581,603]" gridcol="10" gridrow="5" pageId="48" pageNumber="49">59%</td>
|
||
</tr>
|
||
<tr box="[108,1525,619,641]" gridrow="6" pageId="48" pageNumber="49">
|
||
<th box="[108,163,619,641]" gridcol="0" gridrow="6" pageId="48" pageNumber="49">Mx5</th>
|
||
<td box="[212,272,619,641]" gridcol="1" gridrow="6" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,619,641]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,619,641]" gridcol="2" gridrow="6" pageId="48" pageNumber="49">77%</td>
|
||
<td box="[455,522,619,641]" gridcol="3" gridrow="6" pageId="48" pageNumber="49">72%</td>
|
||
<td box="[584,639,619,641]" gridcol="4" gridrow="6" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,619,641]" gridcol="5" gridrow="6" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,619,641]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,619,641]" gridcol="6" gridrow="6" pageId="48" pageNumber="49">74.5%</td>
|
||
<td box="[924,1019,619,641]" gridcol="7" gridrow="6" pageId="48" pageNumber="49">50%,?</td>
|
||
<td box="[1073,1172,619,641]" gridcol="8" gridrow="6" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,619,641]" gridcol="9" gridrow="6" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,619,641]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,619,641]" gridcol="10" gridrow="6" pageId="48" pageNumber="49">56%</td>
|
||
</tr>
|
||
<tr box="[108,1525,656,679]" gridrow="7" pageId="48" pageNumber="49">
|
||
<th box="[108,163,656,679]" gridcol="0" gridrow="7" pageId="48" pageNumber="49">Mx6</th>
|
||
<td box="[212,272,656,679]" gridcol="1" gridrow="7" pageId="48" pageNumber="49">–</td>
|
||
<td box="[326,392,656,679]" gridcol="2" gridrow="7" pageId="48" pageNumber="49">
|
||
<emphasis box="[326,338,657,679]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[455,522,656,679]" gridcol="3" gridrow="7" pageId="48" pageNumber="49">71%</td>
|
||
<td box="[584,639,656,679]" gridcol="4" gridrow="7" pageId="48" pageNumber="49">85%</td>
|
||
<td box="[698,750,656,679]" gridcol="5" gridrow="7" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,657,679]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,656,679]" gridcol="6" gridrow="7" pageId="48" pageNumber="49">78%</td>
|
||
<td box="[924,1019,656,679]" gridcol="7" gridrow="7" pageId="48" pageNumber="49">50%, 56%</td>
|
||
<td box="[1073,1172,656,679]" gridcol="8" gridrow="7" pageId="48" pageNumber="49">74%</td>
|
||
<td box="[1224,1327,656,679]" gridcol="9" gridrow="7" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,657,679]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,656,679]" gridcol="10" gridrow="7" pageId="48" pageNumber="49">63%</td>
|
||
</tr>
|
||
<tr box="[108,1525,694,716]" gridrow="8" pageId="48" pageNumber="49">
|
||
<th box="[108,163,694,716]" gridcol="0" gridrow="8" pageId="48" pageNumber="49">Mx7</th>
|
||
<td box="[212,272,694,716]" gridcol="1" gridrow="8" pageId="48" pageNumber="49">74%</td>
|
||
<td box="[326,392,694,716]" gridcol="2" gridrow="8" pageId="48" pageNumber="49">81%</td>
|
||
<td box="[455,522,694,716]" gridcol="3" gridrow="8" pageId="48" pageNumber="49">
|
||
<emphasis box="[455,467,694,716]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[584,639,694,716]" gridcol="4" gridrow="8" pageId="48" pageNumber="49">85%</td>
|
||
<td box="[698,750,694,716]" gridcol="5" gridrow="8" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,694,716]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,694,716]" gridcol="6" gridrow="8" pageId="48" pageNumber="49">80%</td>
|
||
<td box="[924,1019,694,716]" gridcol="7" gridrow="8" pageId="48" pageNumber="49">45%,?</td>
|
||
<td box="[1073,1172,694,716]" gridcol="8" gridrow="8" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,694,716]" gridcol="9" gridrow="8" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,694,716]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,694,716]" gridcol="10" gridrow="8" pageId="48" pageNumber="49">63%</td>
|
||
</tr>
|
||
<tr box="[108,1525,732,754]" gridrow="9" pageId="48" pageNumber="49">
|
||
<th box="[108,163,732,754]" gridcol="0" gridrow="9" pageId="48" pageNumber="49">Mx8</th>
|
||
<td box="[212,272,732,754]" gridcol="1" gridrow="9" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[326,392,732,754]" gridcol="2" gridrow="9" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,732,754]" gridcol="3" gridrow="9" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,732,754]" gridcol="4" gridrow="9" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,732,754]" gridcol="5" gridrow="9" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,732,754]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,732,754]" gridcol="6" gridrow="9" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[924,1019,732,754]" gridcol="7" gridrow="9" pageId="48" pageNumber="49">53%, 51%</td>
|
||
<td box="[1073,1172,732,754]" gridcol="8" gridrow="9" pageId="48" pageNumber="49">69%</td>
|
||
<td box="[1224,1327,732,754]" gridcol="9" gridrow="9" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,732,754]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,732,754]" gridcol="10" gridrow="9" pageId="48" pageNumber="49">61%</td>
|
||
</tr>
|
||
<tr box="[108,1525,769,791]" gridrow="10" pageId="48" pageNumber="49">
|
||
<th box="[108,163,769,791]" gridcol="0" gridrow="10" pageId="48" pageNumber="49">Mx9</th>
|
||
<td box="[212,272,769,791]" gridcol="1" gridrow="10" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[326,392,769,791]" gridcol="2" gridrow="10" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,769,791]" gridcol="3" gridrow="10" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,769,791]" gridcol="4" gridrow="10" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,769,791]" gridcol="5" gridrow="10" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,769,791]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,769,791]" gridcol="6" gridrow="10" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[924,1019,769,791]" gridcol="7" gridrow="10" pageId="48" pageNumber="49">54%,?</td>
|
||
<td box="[1073,1172,769,791]" gridcol="8" gridrow="10" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,769,791]" gridcol="9" gridrow="10" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,769,791]" gridcol="10" gridrow="10" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,769,791]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,807,829]" gridrow="11" pageId="48" pageNumber="49">
|
||
<th box="[108,163,807,829]" gridcol="0" gridrow="11" pageId="48" pageNumber="49">Mx10</th>
|
||
<td box="[212,272,807,829]" gridcol="1" gridrow="11" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[326,392,807,829]" gridcol="2" gridrow="11" pageId="48" pageNumber="49">72%</td>
|
||
<td box="[455,522,807,829]" gridcol="3" gridrow="11" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,807,829]" gridcol="4" gridrow="11" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,807,829]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,807,829]" gridcol="5" gridrow="11" pageId="48" pageNumber="49">77%</td>
|
||
<td box="[811,870,807,829]" gridcol="6" gridrow="11" pageId="48" pageNumber="49">77.7%</td>
|
||
<td box="[924,1019,807,829]" gridcol="7" gridrow="11" pageId="48" pageNumber="49">?, 54%</td>
|
||
<td box="[1073,1172,807,829]" gridcol="8" gridrow="11" pageId="48" pageNumber="49">73%</td>
|
||
<td box="[1224,1327,807,829]" gridcol="9" gridrow="11" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,807,829]" gridcol="10" gridrow="11" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,807,829]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,844,867]" gridrow="12" pageId="48" pageNumber="49">
|
||
<th box="[108,163,844,867]" gridcol="0" gridrow="12" pageId="48" pageNumber="49">Mx11</th>
|
||
<td box="[212,272,844,867]" gridcol="1" gridrow="12" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[326,392,844,867]" gridcol="2" gridrow="12" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,844,867]" gridcol="3" gridrow="12" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,844,867]" gridcol="4" gridrow="12" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,844,867]" gridcol="5" gridrow="12" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,845,867]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,844,867]" gridcol="6" gridrow="12" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[924,1019,844,867]" gridcol="7" gridrow="12" pageId="48" pageNumber="49">?, 50%</td>
|
||
<td box="[1073,1172,844,867]" gridcol="8" gridrow="12" pageId="48" pageNumber="49">72%</td>
|
||
<td box="[1224,1327,844,867]" gridcol="9" gridrow="12" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,844,867]" gridcol="10" gridrow="12" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,845,867]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,882,904]" gridrow="13" pageId="48" pageNumber="49">
|
||
<th box="[108,163,882,904]" gridcol="0" gridrow="13" pageId="48" pageNumber="49">Mx12</th>
|
||
<td box="[212,272,882,904]" gridcol="1" gridrow="13" pageId="48" pageNumber="49">–</td>
|
||
<td box="[326,392,882,904]" gridcol="2" gridrow="13" pageId="48" pageNumber="49">
|
||
<emphasis box="[326,338,882,904]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[455,522,882,904]" gridcol="3" gridrow="13" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,882,904]" gridcol="4" gridrow="13" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,882,904]" gridcol="5" gridrow="13" pageId="48" pageNumber="49">–</td>
|
||
<td box="[811,870,882,904]" gridcol="6" gridrow="13" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,882,904]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,882,904]" gridcol="7" gridrow="13" pageId="48" pageNumber="49">47%,?</td>
|
||
<td box="[1073,1172,882,904]" gridcol="8" gridrow="13" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,882,904]" gridcol="9" gridrow="13" pageId="48" pageNumber="49">
|
||
<emphasis box="[1224,1236,882,904]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[1376,1525,882,904]" gridcol="10" gridrow="13" pageId="48" pageNumber="49">63%</td>
|
||
</tr>
|
||
<tr box="[108,1525,920,942]" gridrow="14" pageId="48" pageNumber="49">
|
||
<th box="[108,163,920,942]" gridcol="0" gridrow="14" pageId="48" pageNumber="49">Mx13</th>
|
||
<td box="[212,272,920,942]" gridcol="1" gridrow="14" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,920,942]" gridcol="2" gridrow="14" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,920,942]" gridcol="3" gridrow="14" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,920,942]" gridcol="4" gridrow="14" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[698,750,920,942]" gridcol="5" gridrow="14" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,920,942]" gridcol="6" gridrow="14" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,920,942]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,920,942]" gridcol="7" gridrow="14" pageId="48" pageNumber="49">49%, 51%</td>
|
||
<td box="[1073,1172,920,942]" gridcol="8" gridrow="14" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,920,942]" gridcol="9" gridrow="14" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,920,942]" gridcol="10" gridrow="14" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,920,942]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,957,979]" gridrow="15" pageId="48" pageNumber="49">
|
||
<th box="[108,163,957,979]" gridcol="0" gridrow="15" pageId="48" pageNumber="49">Mx14</th>
|
||
<td box="[212,272,957,979]" gridcol="1" gridrow="15" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,957,979]" gridcol="2" gridrow="15" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,957,979]" gridcol="3" gridrow="15" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,957,979]" gridcol="4" gridrow="15" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[698,750,957,979]" gridcol="5" gridrow="15" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,957,979]" gridcol="6" gridrow="15" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,957,979]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,957,979]" gridcol="7" gridrow="15" pageId="48" pageNumber="49">50%,?</td>
|
||
<td box="[1073,1172,957,979]" gridcol="8" gridrow="15" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,957,979]" gridcol="9" gridrow="15" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,957,979]" gridcol="10" gridrow="15" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,957,979]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,995,1017]" gridrow="16" pageId="48" pageNumber="49">
|
||
<th box="[108,163,995,1017]" gridcol="0" gridrow="16" pageId="48" pageNumber="49">Mx15</th>
|
||
<td box="[212,272,995,1017]" gridcol="1" gridrow="16" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,995,1017]" gridcol="2" gridrow="16" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,995,1017]" gridcol="3" gridrow="16" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,995,1017]" gridcol="4" gridrow="16" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[698,750,995,1017]" gridcol="5" gridrow="16" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,995,1017]" gridcol="6" gridrow="16" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,995,1017]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,995,1017]" gridcol="7" gridrow="16" pageId="48" pageNumber="49">54%, 53%</td>
|
||
<td box="[1073,1172,995,1017]" gridcol="8" gridrow="16" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,995,1017]" gridcol="9" gridrow="16" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,995,1017]" gridcol="10" gridrow="16" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,995,1017]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1032,1055]" gridrow="17" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1032,1055]" gridcol="0" gridrow="17" pageId="48" pageNumber="49">Mx16</th>
|
||
<td box="[212,272,1032,1055]" gridcol="1" gridrow="17" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,1032,1055]" gridcol="2" gridrow="17" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,1032,1055]" gridcol="3" gridrow="17" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1032,1055]" gridcol="4" gridrow="17" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[698,750,1032,1055]" gridcol="5" gridrow="17" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,1032,1055]" gridcol="6" gridrow="17" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,1033,1055]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,1032,1055]" gridcol="7" gridrow="17" pageId="48" pageNumber="49">?,?</td>
|
||
<td box="[1073,1172,1032,1055]" gridcol="8" gridrow="17" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1032,1055]" gridcol="9" gridrow="17" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1032,1055]" gridcol="10" gridrow="17" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1033,1055]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1070,1121]" gridrow="18" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1070,1121]" gridcol="0" gridrow="18" pageId="48" pageNumber="49">Dn1</th>
|
||
<td box="[212,272,1070,1121]" gridcol="1" gridrow="18" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,1070,1092]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,1070,1121]" gridcol="2" gridrow="18" pageId="48" pageNumber="49">170%</td>
|
||
<td box="[455,522,1070,1121]" gridcol="3" gridrow="18" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1070,1121]" gridcol="4" gridrow="18" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1070,1092]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1070,1121]" gridcol="5" gridrow="18" pageId="48" pageNumber="49">140%</td>
|
||
<td box="[811,870,1070,1121]" gridcol="6" gridrow="18" pageId="48" pageNumber="49">155%</td>
|
||
<td box="[924,1019,1070,1121]" gridcol="7" gridrow="18" pageId="48" pageNumber="49">130%, 120%</td>
|
||
<td box="[1073,1172,1070,1121]" gridcol="8" gridrow="18" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1070,1121]" gridcol="9" gridrow="18" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1070,1121]" gridcol="10" gridrow="18" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1070,1092]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1137,1159]" gridrow="19" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1137,1159]" gridcol="0" gridrow="19" pageId="48" pageNumber="49">Dn2</th>
|
||
<td box="[212,272,1137,1159]" gridcol="1" gridrow="19" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,1137,1159]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,1137,1159]" gridcol="2" gridrow="19" pageId="48" pageNumber="49">110%</td>
|
||
<td box="[455,522,1137,1159]" gridcol="3" gridrow="19" pageId="48" pageNumber="49">73%</td>
|
||
<td box="[584,639,1137,1159]" gridcol="4" gridrow="19" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1137,1159]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1137,1159]" gridcol="5" gridrow="19" pageId="48" pageNumber="49">94%</td>
|
||
<td box="[811,870,1137,1159]" gridcol="6" gridrow="19" pageId="48" pageNumber="49">92.3%</td>
|
||
<td box="[924,1019,1137,1159]" gridcol="7" gridrow="19" pageId="48" pageNumber="49">83%, 84%</td>
|
||
<td box="[1073,1172,1137,1159]" gridcol="8" gridrow="19" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1137,1159]" gridcol="9" gridrow="19" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1137,1159]" gridcol="10" gridrow="19" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1137,1159]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1175,1197]" gridrow="20" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1175,1197]" gridcol="0" gridrow="20" pageId="48" pageNumber="49">Dn3</th>
|
||
<td box="[212,272,1175,1197]" gridcol="1" gridrow="20" pageId="48" pageNumber="49">–</td>
|
||
<td box="[326,392,1175,1197]" gridcol="2" gridrow="20" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,1175,1197]" gridcol="3" gridrow="20" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1175,1197]" gridcol="4" gridrow="20" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1175,1197]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1175,1197]" gridcol="5" gridrow="20" pageId="48" pageNumber="49">88%</td>
|
||
<td box="[811,870,1175,1197]" gridcol="6" gridrow="20" pageId="48" pageNumber="49">88%</td>
|
||
<td box="[924,1019,1175,1197]" gridcol="7" gridrow="20" pageId="48" pageNumber="49">?, 70%</td>
|
||
<td box="[1073,1172,1175,1197]" gridcol="8" gridrow="20" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1175,1197]" gridcol="9" gridrow="20" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1175,1197]" gridcol="10" gridrow="20" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1175,1197]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1212,1234]" gridrow="21" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1212,1234]" gridcol="0" gridrow="21" pageId="48" pageNumber="49">Dn4</th>
|
||
<td box="[212,272,1212,1234]" gridcol="1" gridrow="21" pageId="48" pageNumber="49">82%</td>
|
||
<td box="[326,392,1212,1234]" gridcol="2" gridrow="21" pageId="48" pageNumber="49">87%</td>
|
||
<td box="[455,522,1212,1234]" gridcol="3" gridrow="21" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1212,1234]" gridcol="4" gridrow="21" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1212,1234]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1212,1234]" gridcol="5" gridrow="21" pageId="48" pageNumber="49">84%</td>
|
||
<td box="[811,870,1212,1234]" gridcol="6" gridrow="21" pageId="48" pageNumber="49">84.3%</td>
|
||
<td box="[924,1019,1212,1234]" gridcol="7" gridrow="21" pageId="48" pageNumber="49">64%, 65%</td>
|
||
<td box="[1073,1172,1212,1234]" gridcol="8" gridrow="21" pageId="48" pageNumber="49">78%</td>
|
||
<td box="[1224,1327,1212,1234]" gridcol="9" gridrow="21" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1212,1234]" gridcol="10" gridrow="21" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1212,1234]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1250,1272]" gridrow="22" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1250,1272]" gridcol="0" gridrow="22" pageId="48" pageNumber="49">Dn5</th>
|
||
<td box="[212,272,1250,1272]" gridcol="1" gridrow="22" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,1250,1272]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,1250,1272]" gridcol="2" gridrow="22" pageId="48" pageNumber="49">120%</td>
|
||
<td box="[455,522,1250,1272]" gridcol="3" gridrow="22" pageId="48" pageNumber="49">73%</td>
|
||
<td box="[584,639,1250,1272]" gridcol="4" gridrow="22" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1250,1272]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1250,1272]" gridcol="5" gridrow="22" pageId="48" pageNumber="49">77%</td>
|
||
<td box="[811,870,1250,1272]" gridcol="6" gridrow="22" pageId="48" pageNumber="49">90%</td>
|
||
<td box="[924,1019,1250,1272]" gridcol="7" gridrow="22" pageId="48" pageNumber="49">58%,?</td>
|
||
<td box="[1073,1172,1250,1272]" gridcol="8" gridrow="22" pageId="48" pageNumber="49">70%</td>
|
||
<td box="[1224,1327,1250,1272]" gridcol="9" gridrow="22" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1250,1272]" gridcol="10" gridrow="22" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1250,1272]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1287,1309]" gridrow="23" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1287,1309]" gridcol="0" gridrow="23" pageId="48" pageNumber="49">Dn6</th>
|
||
<td box="[212,272,1287,1309]" gridcol="1" gridrow="23" pageId="48" pageNumber="49">83%</td>
|
||
<td box="[326,392,1287,1309]" gridcol="2" gridrow="23" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,1287,1309]" gridcol="3" gridrow="23" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1287,1309]" gridcol="4" gridrow="23" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,1287,1309]" gridcol="5" gridrow="23" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,1287,1309]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,1287,1309]" gridcol="6" gridrow="23" pageId="48" pageNumber="49">83%</td>
|
||
<td box="[924,1019,1287,1309]" gridcol="7" gridrow="23" pageId="48" pageNumber="49">61%, 58%</td>
|
||
<td box="[1073,1172,1287,1309]" gridcol="8" gridrow="23" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1287,1309]" gridcol="9" gridrow="23" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1287,1309]" gridcol="10" gridrow="23" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1287,1309]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1325,1347]" gridrow="24" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1325,1347]" gridcol="0" gridrow="24" pageId="48" pageNumber="49">Dn7</th>
|
||
<td box="[212,272,1325,1347]" gridcol="1" gridrow="24" pageId="48" pageNumber="49">79%</td>
|
||
<td box="[326,392,1325,1347]" gridcol="2" gridrow="24" pageId="48" pageNumber="49">130%</td>
|
||
<td box="[455,522,1325,1347]" gridcol="3" gridrow="24" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[584,639,1325,1347]" gridcol="4" gridrow="24" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1325,1347]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1325,1347]" gridcol="5" gridrow="24" pageId="48" pageNumber="49">82%</td>
|
||
<td box="[811,870,1325,1347]" gridcol="6" gridrow="24" pageId="48" pageNumber="49">91.8%</td>
|
||
<td box="[924,1019,1325,1347]" gridcol="7" gridrow="24" pageId="48" pageNumber="49">?, 54%</td>
|
||
<td box="[1073,1172,1325,1347]" gridcol="8" gridrow="24" pageId="48" pageNumber="49">90%</td>
|
||
<td box="[1224,1327,1325,1347]" gridcol="9" gridrow="24" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1325,1347]" gridcol="10" gridrow="24" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1325,1347]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1362,1385]" gridrow="25" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1362,1385]" gridcol="0" gridrow="25" pageId="48" pageNumber="49">Dn8</th>
|
||
<td box="[212,272,1362,1385]" gridcol="1" gridrow="25" pageId="48" pageNumber="49">
|
||
<emphasis box="[212,224,1363,1385]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[326,392,1362,1385]" gridcol="2" gridrow="25" pageId="48" pageNumber="49">79%</td>
|
||
<td box="[455,522,1362,1385]" gridcol="3" gridrow="25" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1362,1385]" gridcol="4" gridrow="25" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,1362,1385]" gridcol="5" gridrow="25" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,1363,1385]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,1362,1385]" gridcol="6" gridrow="25" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[924,1019,1362,1385]" gridcol="7" gridrow="25" pageId="48" pageNumber="49">53%,?</td>
|
||
<td box="[1073,1172,1362,1385]" gridcol="8" gridrow="25" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1362,1385]" gridcol="9" gridrow="25" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1362,1385]" gridcol="10" gridrow="25" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1363,1385]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1400,1422]" gridrow="26" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1400,1422]" gridcol="0" gridrow="26" pageId="48" pageNumber="49">Dn9</th>
|
||
<td box="[212,272,1400,1422]" gridcol="1" gridrow="26" pageId="48" pageNumber="49">77%</td>
|
||
<td box="[326,392,1400,1422]" gridcol="2" gridrow="26" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,1400,1422]" gridcol="3" gridrow="26" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1400,1422]" gridcol="4" gridrow="26" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,1400,1422]" gridcol="5" gridrow="26" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,1400,1422]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,1400,1422]" gridcol="6" gridrow="26" pageId="48" pageNumber="49">77%</td>
|
||
<td box="[924,1019,1400,1422]" gridcol="7" gridrow="26" pageId="48" pageNumber="49">53%,?</td>
|
||
<td box="[1073,1172,1400,1422]" gridcol="8" gridrow="26" pageId="48" pageNumber="49">80%</td>
|
||
<td box="[1224,1327,1400,1422]" gridcol="9" gridrow="26" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1400,1422]" gridcol="10" gridrow="26" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1400,1422]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1438,1460]" gridrow="27" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1438,1460]" gridcol="0" gridrow="27" pageId="48" pageNumber="49">Dn10</th>
|
||
<td box="[212,272,1438,1460]" gridcol="1" gridrow="27" pageId="48" pageNumber="49">69%</td>
|
||
<td box="[326,392,1438,1460]" gridcol="2" gridrow="27" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,1438,1460]" gridcol="3" gridrow="27" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1438,1460]" gridcol="4" gridrow="27" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1438,1460]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1438,1460]" gridcol="5" gridrow="27" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[811,870,1438,1460]" gridcol="6" gridrow="27" pageId="48" pageNumber="49">72.5%</td>
|
||
<td box="[924,1019,1438,1460]" gridcol="7" gridrow="27" pageId="48" pageNumber="49">57%,?</td>
|
||
<td box="[1073,1172,1438,1460]" gridcol="8" gridrow="27" pageId="48" pageNumber="49">86%</td>
|
||
<td box="[1224,1327,1438,1460]" gridcol="9" gridrow="27" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1438,1460]" gridcol="10" gridrow="27" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1438,1460]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1475,1497]" gridrow="28" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1475,1497]" gridcol="0" gridrow="28" pageId="48" pageNumber="49">Dn11</th>
|
||
<td box="[212,272,1475,1497]" gridcol="1" gridrow="28" pageId="48" pageNumber="49">69%</td>
|
||
<td box="[326,392,1475,1497]" gridcol="2" gridrow="28" pageId="48" pageNumber="49">–</td>
|
||
<td box="[455,522,1475,1497]" gridcol="3" gridrow="28" pageId="48" pageNumber="49">–</td>
|
||
<td box="[584,639,1475,1497]" gridcol="4" gridrow="28" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,1475,1497]" gridcol="5" gridrow="28" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,1475,1497]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,1475,1497]" gridcol="6" gridrow="28" pageId="48" pageNumber="49">69%</td>
|
||
<td box="[924,1019,1475,1497]" gridcol="7" gridrow="28" pageId="48" pageNumber="49">53%,?</td>
|
||
<td box="[1073,1172,1475,1497]" gridcol="8" gridrow="28" pageId="48" pageNumber="49">74%</td>
|
||
<td box="[1224,1327,1475,1497]" gridcol="9" gridrow="28" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1475,1497]" gridcol="10" gridrow="28" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1475,1497]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1513,1535]" gridrow="29" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1513,1535]" gridcol="0" gridrow="29" pageId="48" pageNumber="49">Dn12</th>
|
||
<td box="[212,272,1513,1535]" gridcol="1" gridrow="29" pageId="48" pageNumber="49">70%</td>
|
||
<td box="[326,392,1513,1535]" gridcol="2" gridrow="29" pageId="48" pageNumber="49">
|
||
<emphasis box="[326,338,1513,1535]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[455,522,1513,1535]" gridcol="3" gridrow="29" pageId="48" pageNumber="49">67%</td>
|
||
<td box="[584,639,1513,1535]" gridcol="4" gridrow="29" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1513,1535]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1513,1535]" gridcol="5" gridrow="29" pageId="48" pageNumber="49">76%</td>
|
||
<td box="[811,870,1513,1535]" gridcol="6" gridrow="29" pageId="48" pageNumber="49">71%</td>
|
||
<td box="[924,1019,1513,1535]" gridcol="7" gridrow="29" pageId="48" pageNumber="49">?</td>
|
||
<td box="[1073,1172,1513,1535]" gridcol="8" gridrow="29" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1513,1535]" gridcol="9" gridrow="29" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1513,1535]" gridcol="10" gridrow="29" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1513,1535]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1550,1573]" gridrow="30" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1550,1573]" gridcol="0" gridrow="30" pageId="48" pageNumber="49">Dn13</th>
|
||
<td box="[212,272,1550,1573]" gridcol="1" gridrow="30" pageId="48" pageNumber="49">80%</td>
|
||
<td box="[326,392,1550,1573]" gridcol="2" gridrow="30" pageId="48" pageNumber="49">60%</td>
|
||
<td box="[455,522,1550,1573]" gridcol="3" gridrow="30" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1550,1573]" gridcol="4" gridrow="30" pageId="48" pageNumber="49">–</td>
|
||
<td box="[698,750,1550,1573]" gridcol="5" gridrow="30" pageId="48" pageNumber="49">
|
||
<emphasis box="[698,710,1551,1573]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[811,870,1550,1573]" gridcol="6" gridrow="30" pageId="48" pageNumber="49">70%</td>
|
||
<td box="[924,1019,1550,1573]" gridcol="7" gridrow="30" pageId="48" pageNumber="49">50%,?</td>
|
||
<td box="[1073,1172,1550,1573]" gridcol="8" gridrow="30" pageId="48" pageNumber="49">75%</td>
|
||
<td box="[1224,1327,1550,1573]" gridcol="9" gridrow="30" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1550,1573]" gridcol="10" gridrow="30" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1551,1573]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1588,1610]" gridrow="31" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1588,1610]" gridcol="0" gridrow="31" pageId="48" pageNumber="49">Dn14</th>
|
||
<td box="[212,272,1588,1610]" gridcol="1" gridrow="31" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,1588,1610]" gridcol="2" gridrow="31" pageId="48" pageNumber="49">79%</td>
|
||
<td box="[455,522,1588,1610]" gridcol="3" gridrow="31" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1588,1610]" gridcol="4" gridrow="31" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1588,1610]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1588,1610]" gridcol="5" gridrow="31" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,1588,1610]" gridcol="6" gridrow="31" pageId="48" pageNumber="49">79%</td>
|
||
<td box="[924,1019,1588,1610]" gridcol="7" gridrow="31" pageId="48" pageNumber="49">50%,?</td>
|
||
<td box="[1073,1172,1588,1610]" gridcol="8" gridrow="31" pageId="48" pageNumber="49">72%</td>
|
||
<td box="[1224,1327,1588,1610]" gridcol="9" gridrow="31" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1588,1610]" gridcol="10" gridrow="31" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1588,1610]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1625,1648]" gridrow="32" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1625,1648]" gridcol="0" gridrow="32" pageId="48" pageNumber="49">Dn15</th>
|
||
<td box="[212,272,1625,1648]" gridcol="1" gridrow="32" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,1625,1648]" gridcol="2" gridrow="32" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,1625,1648]" gridcol="3" gridrow="32" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1625,1648]" gridcol="4" gridrow="32" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1626,1648]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1625,1648]" gridcol="5" gridrow="32" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,1625,1648]" gridcol="6" gridrow="32" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,1626,1648]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,1625,1648]" gridcol="7" gridrow="32" pageId="48" pageNumber="49">58%,?</td>
|
||
<td box="[1073,1172,1625,1648]" gridcol="8" gridrow="32" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1224,1327,1625,1648]" gridcol="9" gridrow="32" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1625,1648]" gridcol="10" gridrow="32" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1626,1648]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1663,1685]" gridrow="33" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1663,1685]" gridcol="0" gridrow="33" pageId="48" pageNumber="49">Dn16</th>
|
||
<td box="[212,272,1663,1685]" gridcol="1" gridrow="33" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,1663,1685]" gridcol="2" gridrow="33" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,1663,1685]" gridcol="3" gridrow="33" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1663,1685]" gridcol="4" gridrow="33" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1663,1685]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1663,1685]" gridcol="5" gridrow="33" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,1663,1685]" gridcol="6" gridrow="33" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,1663,1685]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,1663,1685]" gridcol="7" gridrow="33" pageId="48" pageNumber="49">53%,?</td>
|
||
<td box="[1073,1172,1663,1685]" gridcol="8" gridrow="33" pageId="48" pageNumber="49">83%</td>
|
||
<td box="[1224,1327,1663,1685]" gridcol="9" gridrow="33" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1663,1685]" gridcol="10" gridrow="33" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1663,1685]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1701,1723]" gridrow="34" pageId="48" pageNumber="49">
|
||
<th box="[108,163,1701,1723]" gridcol="0" gridrow="34" pageId="48" pageNumber="49">Dn17</th>
|
||
<td box="[212,272,1701,1723]" gridcol="1" gridrow="34" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[326,392,1701,1723]" gridcol="2" gridrow="34" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[455,522,1701,1723]" gridcol="3" gridrow="34" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[584,639,1701,1723]" gridcol="4" gridrow="34" pageId="48" pageNumber="49">
|
||
<emphasis box="[584,596,1701,1723]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[698,750,1701,1723]" gridcol="5" gridrow="34" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[811,870,1701,1723]" gridcol="6" gridrow="34" pageId="48" pageNumber="49">
|
||
<emphasis box="[811,823,1701,1723]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
<td box="[924,1019,1701,1723]" gridcol="7" gridrow="34" pageId="48" pageNumber="49">65%,?</td>
|
||
<td box="[1073,1172,1701,1723]" gridcol="8" gridrow="34" pageId="48" pageNumber="49">n/a</td>
|
||
<td box="[1224,1327,1701,1723]" gridcol="9" gridrow="34" pageId="48" pageNumber="49">–</td>
|
||
<td box="[1376,1525,1701,1723]" gridcol="10" gridrow="34" pageId="48" pageNumber="49">
|
||
<emphasis box="[1376,1388,1701,1723]" italics="true" pageId="48" pageNumber="49">–</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825000" ID-Zenodo-Dep="5825000" httpUri="https://zenodo.org/record/5825000/files/figure.png" pageId="49" pageNumber="50" startId="49.[524,589,706,728]" targetBox="[506,1454,236,674]" targetPageId="49">
|
||
<paragraph blockId="49.[524,1515,706,874]" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" pageId="49" pageNumber="50">
|
||
Figure 14 Bivariate scatterplot showing the relationship between maxillary tooth count with maturity among 14 specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[871,1060,735,756]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[871,1060,735,756]" italics="true" pageId="49" pageNumber="50">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which maxillary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Maxillary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-14
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFF5FF99FDEBFC3633DDFBB2" httpUri="http://table.plazi.org/id/DF5C84E2FFF5FF99FDEBFC3633DDFBB2" pageId="49" pageNumber="50" startId="49.[525,580,958,979]" targetBox="[516,1508,1058,1668]" targetIsTable="true" targetPageId="49">
|
||
<paragraph blockId="49.[525,1515,957,1038]" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[525,1416,957,979]" pageId="49" pageNumber="50">
|
||
Table 12 Summary of maxillary tooth count data and growth rank in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1225,1411,958,979]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1225,1411,958,979]" italics="true" pageId="49" pageNumber="50">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of raw and ranked data used for the Spearman correlation test between maturity and maxillary tooth count in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[613,669,1017,1038]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[613,669,1017,1038]" italics="true" pageId="49" pageNumber="50">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="49" pageNumber="50">
|
||
<table box="[516,1508,1058,1668]" gridcols="7" gridrows="15" pageId="49" pageNumber="50">
|
||
<tr box="[516,1508,1058,1139]" gridrow="0" pageId="49" pageNumber="50">
|
||
<th box="[516,709,1058,1139]" gridcol="0" gridrow="0" pageId="49" pageNumber="50">Specimen</th>
|
||
<th box="[725,820,1058,1139]" gridcol="1" gridrow="0" pageId="49" pageNumber="50">Maxillary tooth #</th>
|
||
<th box="[854,980,1058,1139]" gridcol="2" gridrow="0" pageId="49" pageNumber="50">Maxillary tooth # rank</th>
|
||
<th box="[1017,1112,1058,1139]" gridcol="3" gridrow="0" pageId="49" pageNumber="50">
|
||
Maxillary tooth
|
||
<emphasis bold="true" box="[1017,1112,1117,1139]" pageId="49" pageNumber="50">midranks</emphasis>
|
||
</th>
|
||
<th box="[1136,1212,1058,1139]" gridcol="4" gridrow="0" pageId="49" pageNumber="50">Growth stage</th>
|
||
<th box="[1238,1341,1058,1139]" gridcol="5" gridrow="0" pageId="49" pageNumber="50">Growth stage rank</th>
|
||
<th box="[1374,1508,1058,1139]" gridcol="6" gridrow="0" pageId="49" pageNumber="50">Growth stage midranks</th>
|
||
</tr>
|
||
<tr box="[516,1508,1158,1180]" gridrow="1" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1158,1180]" gridcol="0" gridrow="1" pageId="49" pageNumber="50">CMNH 741</th>
|
||
<td box="[725,820,1158,1180]" gridcol="1" gridrow="1" pageId="49" pageNumber="50">15</td>
|
||
<td box="[854,980,1158,1180]" gridcol="2" gridrow="1" pageId="49" pageNumber="50">2</td>
|
||
<td box="[1017,1112,1158,1180]" gridcol="3" gridrow="1" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1158,1179]" pageId="49" pageNumber="50">2</emphasis>
|
||
3
|
||
</td>
|
||
<td box="[1238,1341,1158,1180]" gridcol="5" gridrow="1" pageId="49" pageNumber="50">1</td>
|
||
<td box="[1374,1508,1158,1180]" gridcol="6" gridrow="1" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1158,1179]" pageId="49" pageNumber="50">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1196,1218]" gridrow="2" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1196,1218]" gridcol="0" gridrow="2" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,665,1196,1217]" collectionCode="BMRP" pageId="49" pageNumber="50" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1196,1218]" gridcol="1" gridrow="2" pageId="49" pageNumber="50">16</td>
|
||
<td box="[854,980,1196,1218]" gridcol="2" gridrow="2" pageId="49" pageNumber="50">1</td>
|
||
<td box="[1017,1112,1196,1218]" gridcol="3" gridrow="2" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1196,1217]" pageId="49" pageNumber="50">1</emphasis>
|
||
4
|
||
</td>
|
||
<td box="[1238,1341,1196,1218]" gridcol="5" gridrow="2" pageId="49" pageNumber="50">2</td>
|
||
<td box="[1374,1508,1196,1218]" gridcol="6" gridrow="2" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1196,1217]" pageId="49" pageNumber="50">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1233,1255]" gridrow="3" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1233,1255]" gridcol="0" gridrow="3" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,622,1233,1255]" collectionCode="MOR" pageId="49" pageNumber="50" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1233,1255]" gridcol="1" gridrow="3" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1233,1255]" gridcol="2" gridrow="3" pageId="49" pageNumber="50">3</td>
|
||
<td box="[1017,1112,1233,1255]" gridcol="3" gridrow="3" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1233,1254]" pageId="49" pageNumber="50">7</emphasis>
|
||
7
|
||
</td>
|
||
<td box="[1238,1341,1233,1255]" gridcol="5" gridrow="3" pageId="49" pageNumber="50">3</td>
|
||
<td box="[1374,1508,1233,1255]" gridcol="6" gridrow="3" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1233,1254]" pageId="49" pageNumber="50">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1271,1293]" gridrow="4" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1271,1293]" gridcol="0" gridrow="4" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,709,1271,1293]" collectionCode="TMP" pageId="49" pageNumber="50" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1271,1293]" gridcol="1" gridrow="4" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1271,1293]" gridcol="2" gridrow="4" pageId="49" pageNumber="50">4</td>
|
||
<td box="[1017,1112,1271,1293]" gridcol="3" gridrow="4" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1271,1292]" pageId="49" pageNumber="50">7</emphasis>
|
||
8
|
||
</td>
|
||
<td box="[1238,1341,1271,1293]" gridcol="5" gridrow="4" pageId="49" pageNumber="50">4</td>
|
||
<td box="[1374,1508,1271,1293]" gridcol="6" gridrow="4" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1271,1292]" pageId="49" pageNumber="50">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1308,1330]" gridrow="5" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1308,1330]" gridcol="0" gridrow="5" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,656,1309,1330]" collectionCode="UWBM" pageId="49" pageNumber="50" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1308,1330]" gridcol="1" gridrow="5" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1308,1330]" gridcol="2" gridrow="5" pageId="49" pageNumber="50">5</td>
|
||
<td box="[1017,1112,1308,1330]" gridcol="3" gridrow="5" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1309,1330]" pageId="49" pageNumber="50">7</emphasis>
|
||
11
|
||
</td>
|
||
<td box="[1238,1341,1308,1330]" gridcol="5" gridrow="5" pageId="49" pageNumber="50">5</td>
|
||
<td box="[1374,1508,1308,1330]" gridcol="6" gridrow="5" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1309,1330]" pageId="49" pageNumber="50">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1346,1368]" gridrow="6" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1346,1368]" gridcol="0" gridrow="6" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,632,1346,1368]" collectionCode="RSM" pageId="49" pageNumber="50" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1346,1368]" gridcol="1" gridrow="6" pageId="49" pageNumber="50">11</td>
|
||
<td box="[854,980,1346,1368]" gridcol="2" gridrow="6" pageId="49" pageNumber="50">12</td>
|
||
<td box="[1017,1112,1346,1368]" gridcol="3" gridrow="6" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1041,1346,1367]" pageId="49" pageNumber="50">13</emphasis>
|
||
12
|
||
</td>
|
||
<td box="[1238,1341,1346,1368]" gridcol="5" gridrow="6" pageId="49" pageNumber="50">6</td>
|
||
<td box="[1374,1508,1346,1368]" gridcol="6" gridrow="6" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1404,1346,1367]" pageId="49" pageNumber="50">6.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1384,1406]" gridrow="7" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1384,1406]" gridcol="0" gridrow="7" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,641,1384,1405]" collectionCode="SDSM" pageId="49" pageNumber="50" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1384,1406]" gridcol="1" gridrow="7" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1384,1406]" gridcol="2" gridrow="7" pageId="49" pageNumber="50">6</td>
|
||
<td box="[1017,1112,1384,1406]" gridcol="3" gridrow="7" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1384,1405]" pageId="49" pageNumber="50">7</emphasis>
|
||
12
|
||
</td>
|
||
<td box="[1238,1341,1384,1406]" gridcol="5" gridrow="7" pageId="49" pageNumber="50">7</td>
|
||
<td box="[1374,1508,1384,1406]" gridcol="6" gridrow="7" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1404,1384,1405]" pageId="49" pageNumber="50">6.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1421,1443]" gridrow="8" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1421,1443]" gridcol="0" gridrow="8" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,707,1421,1443]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="49" pageNumber="50" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1421,1443]" gridcol="1" gridrow="8" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1421,1443]" gridcol="2" gridrow="8" pageId="49" pageNumber="50">7</td>
|
||
<td box="[1017,1112,1421,1443]" gridcol="3" gridrow="8" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1421,1442]" pageId="49" pageNumber="50">7</emphasis>
|
||
13
|
||
</td>
|
||
<td box="[1238,1341,1421,1443]" gridcol="5" gridrow="8" pageId="49" pageNumber="50">8</td>
|
||
<td box="[1374,1508,1421,1443]" gridcol="6" gridrow="8" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1386,1421,1442]" pageId="49" pageNumber="50">8</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1459,1481]" gridrow="9" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1459,1481]" gridcol="0" gridrow="9" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,610,1459,1481]" collectionCode="MOR" pageId="49" pageNumber="50" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1459,1481]" gridcol="1" gridrow="9" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1459,1481]" gridcol="2" gridrow="9" pageId="49" pageNumber="50">8</td>
|
||
<td box="[1017,1112,1459,1481]" gridcol="3" gridrow="9" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1459,1480]" pageId="49" pageNumber="50">7</emphasis>
|
||
16
|
||
</td>
|
||
<td box="[1238,1341,1459,1481]" gridcol="5" gridrow="9" pageId="49" pageNumber="50">9</td>
|
||
<td box="[1374,1508,1459,1481]" gridcol="6" gridrow="9" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1404,1459,1480]" pageId="49" pageNumber="50">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1496,1518]" gridrow="10" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1496,1518]" gridcol="0" gridrow="10" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,610,1497,1518]" collectionCode="MOR" pageId="49" pageNumber="50" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1496,1518]" gridcol="1" gridrow="10" pageId="49" pageNumber="50">11</td>
|
||
<td box="[854,980,1496,1518]" gridcol="2" gridrow="10" pageId="49" pageNumber="50">13</td>
|
||
<td box="[1017,1112,1496,1518]" gridcol="3" gridrow="10" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1041,1496,1517]" pageId="49" pageNumber="50">13</emphasis>
|
||
16
|
||
</td>
|
||
<td box="[1238,1341,1496,1518]" gridcol="5" gridrow="10" pageId="49" pageNumber="50">10</td>
|
||
<td box="[1374,1508,1496,1518]" gridcol="6" gridrow="10" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1404,1496,1517]" pageId="49" pageNumber="50">9.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1534,1555]" gridrow="11" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1534,1555]" gridcol="0" gridrow="11" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,646,1534,1555]" collectionCode="LACM" pageId="49" pageNumber="50" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1534,1555]" gridcol="1" gridrow="11" pageId="49" pageNumber="50">11</td>
|
||
<td box="[854,980,1534,1555]" gridcol="2" gridrow="11" pageId="49" pageNumber="50">14</td>
|
||
<td box="[1017,1112,1534,1555]" gridcol="3" gridrow="11" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1041,1534,1555]" pageId="49" pageNumber="50">13</emphasis>
|
||
17
|
||
</td>
|
||
<td box="[1238,1341,1534,1555]" gridcol="5" gridrow="11" pageId="49" pageNumber="50">11</td>
|
||
<td box="[1374,1508,1534,1555]" gridcol="6" gridrow="11" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1416,1534,1555]" pageId="49" pageNumber="50">11.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1572,1593]" gridrow="12" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1572,1593]" gridcol="0" gridrow="12" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,606,1572,1593]" collectionCode="CM" pageId="49" pageNumber="50" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1572,1593]" gridcol="1" gridrow="12" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1572,1593]" gridcol="2" gridrow="12" pageId="49" pageNumber="50">9</td>
|
||
<td box="[1017,1112,1572,1593]" gridcol="3" gridrow="12" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1572,1593]" pageId="49" pageNumber="50">7</emphasis>
|
||
17
|
||
</td>
|
||
<td box="[1238,1341,1572,1593]" gridcol="5" gridrow="12" pageId="49" pageNumber="50">12</td>
|
||
<td box="[1374,1508,1572,1593]" gridcol="6" gridrow="12" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1416,1572,1593]" pageId="49" pageNumber="50">11.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1609,1630]" gridrow="13" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1609,1630]" gridcol="0" gridrow="13" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,610,1609,1630]" collectionCode="MOR" pageId="49" pageNumber="50" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1609,1630]" gridcol="1" gridrow="13" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1609,1630]" gridcol="2" gridrow="13" pageId="49" pageNumber="50">10</td>
|
||
<td box="[1017,1112,1609,1630]" gridcol="3" gridrow="13" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1609,1630]" pageId="49" pageNumber="50">7</emphasis>
|
||
18
|
||
</td>
|
||
<td box="[1238,1341,1609,1630]" gridcol="5" gridrow="13" pageId="49" pageNumber="50">13</td>
|
||
<td box="[1374,1508,1609,1630]" gridcol="6" gridrow="13" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1398,1609,1630]" pageId="49" pageNumber="50">13</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1508,1647,1668]" gridrow="14" pageId="49" pageNumber="50" rowspan-4="1">
|
||
<th box="[516,709,1647,1668]" gridcol="0" gridrow="14" pageId="49" pageNumber="50">
|
||
<materialsCitation box="[516,667,1647,1668]" collectionCode="FMNH" pageId="49" pageNumber="50" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[725,820,1647,1668]" gridcol="1" gridrow="14" pageId="49" pageNumber="50">12</td>
|
||
<td box="[854,980,1647,1668]" gridcol="2" gridrow="14" pageId="49" pageNumber="50">11</td>
|
||
<td box="[1017,1112,1647,1668]" gridcol="3" gridrow="14" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1017,1029,1647,1668]" pageId="49" pageNumber="50">7</emphasis>
|
||
20
|
||
</td>
|
||
<td box="[1238,1341,1647,1668]" gridcol="5" gridrow="14" pageId="49" pageNumber="50">14</td>
|
||
<td box="[1374,1508,1647,1668]" gridcol="6" gridrow="14" pageId="49" pageNumber="50">
|
||
<emphasis bold="true" box="[1374,1398,1647,1668]" pageId="49" pageNumber="50">14</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="49.[498,1542,1752,1900]" pageId="49" pageNumber="50">
|
||
A Shapiro
|
||
<emphasis box="[649,664,1753,1778]" italics="true" pageId="49" pageNumber="50">–</emphasis>
|
||
Wilk test found that the maxillary tooth rank data are not normally distributed (
|
||
<emphasis box="[642,656,1792,1817]" italics="true" pageId="49" pageNumber="50">p</emphasis>
|
||
= 0.004), whereas the growth rank data are normally distributed (
|
||
<emphasis box="[1419,1433,1792,1817]" italics="true" pageId="49" pageNumber="50">p</emphasis>
|
||
= 0.725); a Spearman rank correlation test of growth rank and tooth count ranks was done, which resulted in a statistically nonsignificant (
|
||
<emphasis box="[975,989,1872,1897]" italics="true" pageId="49" pageNumber="50">p</emphasis>
|
||
= 0.073) correlation coefficient (
|
||
<emphasis box="[1379,1390,1872,1897]" italics="true" pageId="49" pageNumber="50">
|
||
<emphasis box="[1390,1400,1884,1900]" italics="true" pageId="49" pageNumber="50">
|
||
<subScript attach="left" box="[1390,1400,1884,1900]" fontSize="7" pageId="49" pageNumber="50">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.494).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825002" ID-Zenodo-Dep="5825002" httpUri="https://zenodo.org/record/5825002/files/figure.png" pageId="50" pageNumber="51" startId="50.[524,589,692,714]" targetBox="[498,1541,249,679]" targetPageId="50">
|
||
<paragraph blockId="50.[524,1516,692,860]" pageId="50" pageNumber="51">
|
||
<emphasis bold="true" pageId="50" pageNumber="51">
|
||
Figure 15 Bivariate scatterplot showing the relationship between dentary tooth count with maturity among 16 specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[762,947,721,742]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[762,947,721,742]" italics="true" pageId="50" pageNumber="51">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which dentary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Dentary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-15
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="50.[498,1542,915,1340]" pageId="50" pageNumber="51">Therefore, a general trend of decrease in tooth count is not seen across the entire data set. This result must be regarded with the caveat that there is a gap in the data, where maxillary tooth counts are currently unknown for subadult specimens. It is possible that tooth counts from future discoveries of subadults will bridge the gap between the extremes and result in a significant value.</paragraph>
|
||
<paragraph blockId="50.[498,1542,915,1340]" pageId="50" pageNumber="51">
|
||
In addition to that, the hypothesis that tooth count is a reliable proxy for maturity from young adults to senescent adults was tested by limiting the statistical test to the corresponding portion of the data set (growth stage ranks 3
|
||
<emphasis box="[1202,1217,1195,1220]" italics="true" pageId="50" pageNumber="51">–</emphasis>
|
||
14); a statistically nonsignificant result was obtained (
|
||
<emphasis box="[919,930,1235,1260]" italics="true" pageId="50" pageNumber="51">
|
||
<emphasis box="[930,940,1246,1262]" italics="true" pageId="50" pageNumber="51">
|
||
<subScript attach="left" box="[930,940,1246,1262]" fontSize="7" pageId="50" pageNumber="51">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.112,
|
||
<emphasis box="[1052,1066,1235,1260]" italics="true" pageId="50" pageNumber="51">p</emphasis>
|
||
= 0.729), indicating that the null hypothesis of no difference could not be rejected. Ergo, maxillary tooth count is an unreliable proxy for maturity among adult-sized specimens.
|
||
</paragraph>
|
||
<paragraph blockId="50.[498,1542,1379,1887]" box="[498,998,1379,1407]" pageId="50" pageNumber="51">
|
||
<heading bold="true" box="[498,998,1379,1407]" fontSize="11" level="3" pageId="50" pageNumber="51" reason="6">
|
||
<emphasis bold="true" box="[498,998,1379,1407]" italics="true" pageId="50" pageNumber="51">Correlation with dentary tooth count</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="50.[498,1542,1379,1887]" pageId="50" pageNumber="51">
|
||
As for the maxilla, an initial increase (from 16 to 17) followed by a decrease in tooth count (from 17 to 14) is seen in the dentary, after which variation is seen (from 14 to 12) among adults (
|
||
<figureCitation box="[676,755,1502,1529]" captionStart="Figure 15" captionStartId="50.[524,589,692,714]" captionTargetBox="[498,1541,249,679]" captionTargetId="figure-357@50.[498,1458,234,664]" captionTargetPageId="50" captionText="Figure 15 Bivariate scatterplot showing the relationship between dentary tooth count with maturity among 16 specimens of Tyrannosaurus rex. Growth rank increases away from the origin (i.e., maturity increases to the right) and corresponds to growth stages for which dentary tooth count was available for a given specimen; that is, the rank does not correspond to growth stage. Dentary tooth rank corresponds to relative tooth count, where low ranks correspond to high tooth counts and low ranks correspond to high tooth counts. Full-size DOI: 10.7717/peerj.9192/fig-15" figureDoi="http://doi.org/10.5281/zenodo.5825002" httpUri="https://zenodo.org/record/5825002/files/figure.png" pageId="50" pageNumber="51">Fig. 15</figureCitation>
|
||
). The raw data were converted to ranks (
|
||
<tableCitation box="[1248,1349,1502,1528]" captionStart="Table 13" captionStartId="51.[525,580,248,269]" captionTargetBox="[516,1501,349,1043]" captionText="Table 13 Summary of dentary tooth count data and growth rank in Tyrannosaurus rex. Summary of raw and ranked data used in the Spearman rank correlation test of maturity and dentary tooth count in T. rex. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FFF7FF9BFDEBFF703375FEF4" pageId="50" pageNumber="51" tableUuid="DF5C84E2FFF7FF9BFDEBFF703375FEF4">Table 13</tableCitation>
|
||
), and a Shapiro
|
||
<emphasis box="[588,603,1543,1568]" italics="true" pageId="50" pageNumber="51">–</emphasis>
|
||
Wilk test of normality found that the dentary tooth count rank data are not normally distributed (
|
||
<emphasis box="[758,772,1583,1608]" italics="true" pageId="50" pageNumber="51">p</emphasis>
|
||
= 0.021), whereas the growth rank data are normally distributed (
|
||
<emphasis box="[508,522,1623,1648]" italics="true" pageId="50" pageNumber="51">p</emphasis>
|
||
= 0.616); a Spearman correlation test of the data resulted a significant (
|
||
<emphasis box="[1373,1387,1623,1648]" italics="true" pageId="50" pageNumber="51">p</emphasis>
|
||
= 0.005) correlation (
|
||
<emphasis box="[644,655,1662,1687]" italics="true" pageId="50" pageNumber="51">
|
||
<emphasis box="[655,665,1674,1690]" italics="true" pageId="50" pageNumber="51">
|
||
<subScript attach="left" box="[655,665,1674,1690]" fontSize="7" pageId="50" pageNumber="51">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.648), indicating a trend of decrease in tooth count across the data set. Among adults, the hypothesis of dentary tooth count as a proxy for maturity was tested by obtaining the correlation coefficient from young adults to senescent adults (from growth rank 3
|
||
<emphasis box="[663,678,1782,1807]" italics="true" pageId="50" pageNumber="51">–</emphasis>
|
||
17), which resulted in a nonsignificant (
|
||
<emphasis box="[1141,1155,1782,1807]" italics="true" pageId="50" pageNumber="51">p</emphasis>
|
||
= 0.074) correlation (
|
||
<emphasis box="[1411,1422,1782,1807]" italics="true" pageId="50" pageNumber="51">
|
||
<emphasis box="[1422,1432,1794,1810]" italics="true" pageId="50" pageNumber="51">
|
||
<subScript attach="left" box="[1422,1432,1794,1810]" fontSize="7" pageId="50" pageNumber="51">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.475), indicating that dentary tooth count is not a reliable proxy for estimating maturity among adult-sized animals.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFF7FF9BFDEBFF703375FEF4" httpUri="http://table.plazi.org/id/DF5C84E2FFF7FF9BFDEBFF703375FEF4" pageId="51" pageNumber="52" startId="51.[525,580,248,269]" targetBox="[516,1501,349,1043]" targetIsTable="true" targetPageId="51">
|
||
<paragraph blockId="51.[525,1515,248,329]" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[525,1392,248,270]" pageId="51" pageNumber="52">
|
||
Table 13 Summary of dentary tooth count data and growth rank in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1201,1386,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1201,1386,249,270]" italics="true" pageId="51" pageNumber="52">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of raw and ranked data used in the Spearman rank correlation test of maturity and dentary tooth count in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[525,581,307,328]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[525,581,307,328]" italics="true" pageId="51" pageNumber="52">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="51" pageNumber="52">
|
||
<table box="[516,1501,349,1043]" gridcols="7" gridrows="18" pageId="51" pageNumber="52">
|
||
<tr box="[516,1501,349,400]" gridrow="0" pageId="51" pageNumber="52">
|
||
<th box="[516,709,349,400]" gridcol="0" gridrow="0" pageId="51" pageNumber="52">Specimen</th>
|
||
<th box="[725,806,349,400]" gridcol="1" gridrow="0" pageId="51" pageNumber="52">Dentary tooth #</th>
|
||
<th box="[848,991,349,400]" gridcol="2" gridrow="0" pageId="51" pageNumber="52">Dentary tooth # rank</th>
|
||
<th box="[1007,1168,349,400]" gridcol="3" gridrow="0" pageId="51" pageNumber="52">Dentary tooth # midranks</th>
|
||
<th box="[1199,1275,349,400]" gridcol="4" gridrow="0" pageId="51" pageNumber="52">Growth stage</th>
|
||
<th box="[1303,1379,349,400]" gridcol="5" gridrow="0" pageId="51" pageNumber="52">Growth rank</th>
|
||
<th box="[1406,1501,349,400]" gridcol="6" gridrow="0" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1501,349,400]" pageId="51" pageNumber="52">Growth midranks</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1501,419,441]" gridrow="1" pageId="51" pageNumber="52">
|
||
<th box="[516,709,419,441]" gridcol="0" gridrow="1" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,642,419,441]" collectionCode="CMNH" pageId="51" pageNumber="52" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,419,441]" gridcol="1" gridrow="1" pageId="51" pageNumber="52">16</td>
|
||
<td box="[848,991,419,441]" gridcol="2" gridrow="1" pageId="51" pageNumber="52">2</td>
|
||
<td box="[1007,1168,419,441]" gridcol="3" gridrow="1" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,419,440]" pageId="51" pageNumber="52">2</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,419,441]" gridcol="4" gridrow="1" pageId="51" pageNumber="52">3</td>
|
||
<td box="[1303,1379,419,441]" gridcol="5" gridrow="1" pageId="51" pageNumber="52">1</td>
|
||
<td box="[1406,1501,419,441]" gridcol="6" gridrow="1" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,419,440]" pageId="51" pageNumber="52">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,457,479]" gridrow="2" pageId="51" pageNumber="52">
|
||
<th box="[516,709,457,479]" gridcol="0" gridrow="2" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,665,457,479]" collectionCode="BMRP" pageId="51" pageNumber="52" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,457,479]" gridcol="1" gridrow="2" pageId="51" pageNumber="52">17</td>
|
||
<td box="[848,991,457,479]" gridcol="2" gridrow="2" pageId="51" pageNumber="52">1</td>
|
||
<td box="[1007,1168,457,479]" gridcol="3" gridrow="2" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,457,478]" pageId="51" pageNumber="52">1</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,457,479]" gridcol="4" gridrow="2" pageId="51" pageNumber="52">4</td>
|
||
<td box="[1303,1379,457,479]" gridcol="5" gridrow="2" pageId="51" pageNumber="52">2</td>
|
||
<td box="[1406,1501,457,479]" gridcol="6" gridrow="2" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,457,478]" pageId="51" pageNumber="52">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,494,516]" gridrow="3" pageId="51" pageNumber="52">
|
||
<th box="[516,709,494,516]" gridcol="0" gridrow="3" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,622,494,516]" collectionCode="MOR" pageId="51" pageNumber="52" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,494,516]" gridcol="1" gridrow="3" pageId="51" pageNumber="52">14</td>
|
||
<td box="[848,991,494,516]" gridcol="2" gridrow="3" pageId="51" pageNumber="52">3</td>
|
||
<td box="[1007,1168,494,516]" gridcol="3" gridrow="3" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,495,516]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,494,516]" gridcol="4" gridrow="3" pageId="51" pageNumber="52">7</td>
|
||
<td box="[1303,1379,494,516]" gridcol="5" gridrow="3" pageId="51" pageNumber="52">3</td>
|
||
<td box="[1406,1501,494,516]" gridcol="6" gridrow="3" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,494,515]" pageId="51" pageNumber="52">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,532,554]" gridrow="4" pageId="51" pageNumber="52">
|
||
<th box="[516,709,532,554]" gridcol="0" gridrow="4" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,709,532,554]" collectionCode="TMP" pageId="51" pageNumber="52" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,532,554]" gridcol="1" gridrow="4" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,532,554]" gridcol="2" gridrow="4" pageId="51" pageNumber="52">8</td>
|
||
<td box="[1007,1168,532,554]" gridcol="3" gridrow="4" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,532,553]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,532,554]" gridcol="4" gridrow="4" pageId="51" pageNumber="52">8</td>
|
||
<td box="[1303,1379,532,554]" gridcol="5" gridrow="4" pageId="51" pageNumber="52">4</td>
|
||
<td box="[1406,1501,532,554]" gridcol="6" gridrow="4" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,532,553]" pageId="51" pageNumber="52">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,570,592]" gridrow="5" pageId="51" pageNumber="52">
|
||
<th box="[516,709,570,592]" gridcol="0" gridrow="5" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,658,570,592]" collectionCode="LACM" pageId="51" pageNumber="52" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,570,592]" gridcol="1" gridrow="5" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,570,592]" gridcol="2" gridrow="5" pageId="51" pageNumber="52">9</td>
|
||
<td box="[1007,1168,570,592]" gridcol="3" gridrow="5" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,570,591]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,570,592]" gridcol="4" gridrow="5" pageId="51" pageNumber="52">9</td>
|
||
<td box="[1303,1379,570,592]" gridcol="5" gridrow="5" pageId="51" pageNumber="52">5</td>
|
||
<td box="[1406,1501,570,592]" gridcol="6" gridrow="5" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,570,591]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,607,629]" gridrow="6" pageId="51" pageNumber="52">
|
||
<th box="[516,709,607,629]" gridcol="0" gridrow="6" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,656,608,629]" collectionCode="UWBM" pageId="51" pageNumber="52" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,607,629]" gridcol="1" gridrow="6" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,607,629]" gridcol="2" gridrow="6" pageId="51" pageNumber="52">10</td>
|
||
<td box="[1007,1168,607,629]" gridcol="3" gridrow="6" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,607,628]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,607,629]" gridcol="4" gridrow="6" pageId="51" pageNumber="52">11</td>
|
||
<td box="[1303,1379,607,629]" gridcol="5" gridrow="6" pageId="51" pageNumber="52">6</td>
|
||
<td box="[1406,1501,607,629]" gridcol="6" gridrow="6" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,607,628]" pageId="51" pageNumber="52">6</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,645,667]" gridrow="7" pageId="51" pageNumber="52">
|
||
<th box="[516,709,645,667]" gridcol="0" gridrow="7" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,632,645,667]" collectionCode="RSM" pageId="51" pageNumber="52" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,645,667]" gridcol="1" gridrow="7" pageId="51" pageNumber="52">14</td>
|
||
<td box="[848,991,645,667]" gridcol="2" gridrow="7" pageId="51" pageNumber="52">4</td>
|
||
<td box="[1007,1168,645,667]" gridcol="3" gridrow="7" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,645,666]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,645,667]" gridcol="4" gridrow="7" pageId="51" pageNumber="52">12</td>
|
||
<td box="[1303,1379,645,667]" gridcol="5" gridrow="7" pageId="51" pageNumber="52">7</td>
|
||
<td box="[1406,1501,645,667]" gridcol="6" gridrow="7" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1436,645,666]" pageId="51" pageNumber="52">7.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,683,705]" gridrow="8" pageId="51" pageNumber="52">
|
||
<th box="[516,709,683,705]" gridcol="0" gridrow="8" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,641,683,704]" collectionCode="SDSM" pageId="51" pageNumber="52" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,683,705]" gridcol="1" gridrow="8" pageId="51" pageNumber="52">14</td>
|
||
<td box="[848,991,683,705]" gridcol="2" gridrow="8" pageId="51" pageNumber="52">5</td>
|
||
<td box="[1007,1168,683,705]" gridcol="3" gridrow="8" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,683,704]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,683,705]" gridcol="4" gridrow="8" pageId="51" pageNumber="52">12</td>
|
||
<td box="[1303,1379,683,705]" gridcol="5" gridrow="8" pageId="51" pageNumber="52">8</td>
|
||
<td box="[1406,1501,683,705]" gridcol="6" gridrow="8" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1436,683,704]" pageId="51" pageNumber="52">7.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,720,742]" gridrow="9" pageId="51" pageNumber="52">
|
||
<th box="[516,709,720,742]" gridcol="0" gridrow="9" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,707,720,742]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="51" pageNumber="52" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,720,742]" gridcol="1" gridrow="9" pageId="51" pageNumber="52">14</td>
|
||
<td box="[848,991,720,742]" gridcol="2" gridrow="9" pageId="51" pageNumber="52">6</td>
|
||
<td box="[1007,1168,720,742]" gridcol="3" gridrow="9" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,720,741]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,720,742]" gridcol="4" gridrow="9" pageId="51" pageNumber="52">13</td>
|
||
<td box="[1303,1379,720,742]" gridcol="5" gridrow="9" pageId="51" pageNumber="52">9</td>
|
||
<td box="[1406,1501,720,742]" gridcol="6" gridrow="9" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1418,720,741]" pageId="51" pageNumber="52">9</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,758,779]" gridrow="10" pageId="51" pageNumber="52">
|
||
<th box="[516,709,758,779]" gridcol="0" gridrow="10" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,674,758,779]" collectionCode="NHMUK" pageId="51" pageNumber="52" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,758,779]" gridcol="1" gridrow="10" pageId="51" pageNumber="52">14</td>
|
||
<td box="[848,991,758,779]" gridcol="2" gridrow="10" pageId="51" pageNumber="52">7</td>
|
||
<td box="[1007,1168,758,779]" gridcol="3" gridrow="10" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1019,758,779]" pageId="51" pageNumber="52">5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,758,779]" gridcol="4" gridrow="10" pageId="51" pageNumber="52">14</td>
|
||
<td box="[1303,1379,758,779]" gridcol="5" gridrow="10" pageId="51" pageNumber="52">10</td>
|
||
<td box="[1406,1501,758,779]" gridcol="6" gridrow="10" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1430,758,779]" pageId="51" pageNumber="52">10</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,795,817]" gridrow="11" pageId="51" pageNumber="52">
|
||
<th box="[516,709,795,817]" gridcol="0" gridrow="11" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,686,795,817]" collectionCode="NMMNH" pageId="51" pageNumber="52" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,795,817]" gridcol="1" gridrow="11" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,795,817]" gridcol="2" gridrow="11" pageId="51" pageNumber="52">11</td>
|
||
<td box="[1007,1168,795,817]" gridcol="3" gridrow="11" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,795,816]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,795,817]" gridcol="4" gridrow="11" pageId="51" pageNumber="52">15</td>
|
||
<td box="[1303,1379,795,817]" gridcol="5" gridrow="11" pageId="51" pageNumber="52">11</td>
|
||
<td box="[1406,1501,795,817]" gridcol="6" gridrow="11" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1430,795,816]" pageId="51" pageNumber="52">11</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,833,855]" gridrow="12" pageId="51" pageNumber="52">
|
||
<th box="[516,709,833,855]" gridcol="0" gridrow="12" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,610,833,855]" collectionCode="MOR" pageId="51" pageNumber="52" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,833,855]" gridcol="1" gridrow="12" pageId="51" pageNumber="52">12</td>
|
||
<td box="[848,991,833,855]" gridcol="2" gridrow="12" pageId="51" pageNumber="52">16</td>
|
||
<td box="[1007,1168,833,855]" gridcol="3" gridrow="12" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,833,854]" pageId="51" pageNumber="52">16.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,833,855]" gridcol="4" gridrow="12" pageId="51" pageNumber="52">16</td>
|
||
<td box="[1303,1379,833,855]" gridcol="5" gridrow="12" pageId="51" pageNumber="52">12</td>
|
||
<td box="[1406,1501,833,855]" gridcol="6" gridrow="12" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1448,833,854]" pageId="51" pageNumber="52">12.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,870,893]" gridrow="13" pageId="51" pageNumber="52">
|
||
<th box="[516,709,870,893]" gridcol="0" gridrow="13" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,610,871,892]" collectionCode="MOR" pageId="51" pageNumber="52" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,870,893]" gridcol="1" gridrow="13" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,870,893]" gridcol="2" gridrow="13" pageId="51" pageNumber="52">12</td>
|
||
<td box="[1007,1168,870,893]" gridcol="3" gridrow="13" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,870,891]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,870,893]" gridcol="4" gridrow="13" pageId="51" pageNumber="52">16</td>
|
||
<td box="[1303,1379,870,893]" gridcol="5" gridrow="13" pageId="51" pageNumber="52">13</td>
|
||
<td box="[1406,1501,870,893]" gridcol="6" gridrow="13" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1448,870,891]" pageId="51" pageNumber="52">12.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,908,929]" gridrow="14" pageId="51" pageNumber="52">
|
||
<th box="[516,709,908,929]" gridcol="0" gridrow="14" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,646,908,929]" collectionCode="LACM" pageId="51" pageNumber="52" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,908,929]" gridcol="1" gridrow="14" pageId="51" pageNumber="52">12</td>
|
||
<td box="[848,991,908,929]" gridcol="2" gridrow="14" pageId="51" pageNumber="52">17</td>
|
||
<td box="[1007,1168,908,929]" gridcol="3" gridrow="14" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,908,929]" pageId="51" pageNumber="52">16.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,908,929]" gridcol="4" gridrow="14" pageId="51" pageNumber="52">17</td>
|
||
<td box="[1303,1379,908,929]" gridcol="5" gridrow="14" pageId="51" pageNumber="52">14</td>
|
||
<td box="[1406,1501,908,929]" gridcol="6" gridrow="14" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1448,908,929]" pageId="51" pageNumber="52">14.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,946,968]" gridrow="15" pageId="51" pageNumber="52">
|
||
<th box="[516,709,946,968]" gridcol="0" gridrow="15" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,606,946,967]" collectionCode="CM" pageId="51" pageNumber="52" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,946,968]" gridcol="1" gridrow="15" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,946,968]" gridcol="2" gridrow="15" pageId="51" pageNumber="52">13</td>
|
||
<td box="[1007,1168,946,968]" gridcol="3" gridrow="15" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,946,967]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,946,968]" gridcol="4" gridrow="15" pageId="51" pageNumber="52">17</td>
|
||
<td box="[1303,1379,946,968]" gridcol="5" gridrow="15" pageId="51" pageNumber="52">15</td>
|
||
<td box="[1406,1501,946,968]" gridcol="6" gridrow="15" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1448,946,967]" pageId="51" pageNumber="52">14.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,983,1005]" gridrow="16" pageId="51" pageNumber="52">
|
||
<th box="[516,709,983,1005]" gridcol="0" gridrow="16" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,610,984,1005]" collectionCode="MOR" pageId="51" pageNumber="52" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,983,1005]" gridcol="1" gridrow="16" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,983,1005]" gridcol="2" gridrow="16" pageId="51" pageNumber="52">14</td>
|
||
<td box="[1007,1168,983,1005]" gridcol="3" gridrow="16" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,983,1004]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,983,1005]" gridcol="4" gridrow="16" pageId="51" pageNumber="52">18</td>
|
||
<td box="[1303,1379,983,1005]" gridcol="5" gridrow="16" pageId="51" pageNumber="52">16</td>
|
||
<td box="[1406,1501,983,1005]" gridcol="6" gridrow="16" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1430,983,1004]" pageId="51" pageNumber="52">16</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1501,1021,1043]" gridrow="17" pageId="51" pageNumber="52">
|
||
<th box="[516,709,1021,1043]" gridcol="0" gridrow="17" pageId="51" pageNumber="52">
|
||
<materialsCitation box="[516,667,1021,1043]" collectionCode="FMNH" pageId="51" pageNumber="52" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[725,806,1021,1043]" gridcol="1" gridrow="17" pageId="51" pageNumber="52">13</td>
|
||
<td box="[848,991,1021,1043]" gridcol="2" gridrow="17" pageId="51" pageNumber="52">15</td>
|
||
<td box="[1007,1168,1021,1043]" gridcol="3" gridrow="17" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1007,1049,1021,1042]" pageId="51" pageNumber="52">11.5</emphasis>
|
||
</td>
|
||
<td box="[1199,1275,1021,1043]" gridcol="4" gridrow="17" pageId="51" pageNumber="52">20</td>
|
||
<td box="[1303,1379,1021,1043]" gridcol="5" gridrow="17" pageId="51" pageNumber="52">17</td>
|
||
<td box="[1406,1501,1021,1043]" gridcol="6" gridrow="17" pageId="51" pageNumber="52">
|
||
<emphasis bold="true" box="[1406,1430,1021,1042]" pageId="51" pageNumber="52">17</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="51.[498,1542,1124,1396]" box="[498,1541,1124,1154]" pageId="51" pageNumber="52">
|
||
<heading bold="true" box="[498,1541,1124,1154]" fontSize="12" level="2" pageId="51" pageNumber="52" reason="0">
|
||
<emphasis bold="true" box="[498,1541,1124,1154]" pageId="51" pageNumber="52">Congruence between chronological age, size, and mass with maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="51.[498,1542,1124,1396]" pageId="51" pageNumber="52">
|
||
Congruence between age, size, and mass (dependent variables) with maturity (independent variable) was tested using Spearman rank correlation. As used here, maturity refers to position of stages along the ontogram. The chronological age of
|
||
<materialsCitation box="[1259,1444,1250,1277]" collectionCode="BMRP" pageId="51" pageNumber="52" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
. is incongruent with maturity, where its chronological age of 15 years is greater than the more mature
|
||
<materialsCitation box="[590,750,1329,1356]" collectionCode="LACM" pageId="51" pageNumber="52" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
that is estimated as 14 years old (
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1154,1388,1330,1356]" journalOrPublisher="Nature" pageId="51" pageNumber="52" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1154,1388,1330,1356]" italics="true" pageId="51" pageNumber="52">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
). Given this discrepancy, the age of
|
||
<materialsCitation box="[775,935,1369,1396]" collectionCode="LACM" pageId="51" pageNumber="52" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
is regarded here as an underestimate.
|
||
</paragraph>
|
||
<paragraph blockId="51.[498,1542,1435,1864]" box="[498,777,1435,1463]" pageId="51" pageNumber="52">
|
||
<heading bold="true" box="[498,777,1435,1463]" fontSize="11" level="3" pageId="51" pageNumber="52" reason="6">
|
||
<emphasis bold="true" box="[498,777,1435,1463]" italics="true" pageId="51" pageNumber="52">Correlation with age</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="51.[498,1542,1435,1864]" pageId="51" pageNumber="52">
|
||
A bivariate scatterplot shows congruence of ranked data between growth rank and age rank (
|
||
<figureCitation box="[571,650,1518,1544]" captionStart="Figure 16" captionStartId="52.[524,589,704,726]" captionTargetBox="[498,1541,234,691]" captionTargetId="figure-336@52.[498,1458,234,677]" captionTargetPageId="52" captionText="Figure 16 Bivariate scatterplot showing the relationship between chronological age with maturity among eight specimens of Tyrannosaurus rex. The comparison is limited to specimens that have been histologically aged; growth stages (x-axis) and chronological age (y-axis) have been converted to ranks. See Table 14 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-16" figureDoi="http://doi.org/10.5281/zenodo.5825004" httpUri="https://zenodo.org/record/5825004/files/figure.png" pageId="51" pageNumber="52">Fig. 16</figureCitation>
|
||
;
|
||
<tableCitation box="[665,766,1518,1544]" captionStart="Table 14" captionStartId="52.[525,580,895,916]" captionTargetBox="[516,1497,996,1351]" captionText="Table 14 Summary of chronological age data and growth rank in Tyrannosaurus rex. Summary of the raw and ranked data used in the Spearman correlation test of maturity and chronological age in T. rex. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FFF0FF9CFDEBFCF73331FC73" pageId="51" pageNumber="52" tableUuid="DF5C84E2FFF0FF9CFDEBFCF73331FC73">Table 14</tableCitation>
|
||
); a Shaprio
|
||
<emphasis box="[903,918,1519,1544]" italics="true" pageId="51" pageNumber="52">–</emphasis>
|
||
Wilk test found that the chronological age ranks and growth ranks are normally distributed (
|
||
<emphasis box="[967,981,1559,1584]" italics="true" pageId="51" pageNumber="52">p</emphasis>
|
||
= 0.885 and 0.933, respectively). A significant (
|
||
<emphasis box="[508,522,1599,1624]" italics="true" pageId="51" pageNumber="52">p</emphasis>
|
||
= 0.000) correlation (
|
||
<emphasis box="[775,786,1599,1624]" italics="true" pageId="51" pageNumber="52">
|
||
<emphasis box="[785,795,1610,1626]" italics="true" pageId="51" pageNumber="52">
|
||
<subScript attach="left" box="[785,795,1610,1626]" fontSize="7" pageId="51" pageNumber="52">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.970) was obtained between age and maturity, indicating that LAG counts are an accurate predictor of maturity. The
|
||
<emphasis box="[1146,1157,1639,1664]" italics="true" pageId="51" pageNumber="52">
|
||
<emphasis box="[1157,1167,1650,1666]" italics="true" pageId="51" pageNumber="52">
|
||
<subScript attach="left" box="[1157,1167,1650,1666]" fontSize="7" pageId="51" pageNumber="52">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
of less than 1.0 is caused by the chronological age of one specimen (
|
||
<materialsCitation box="[922,1085,1677,1704]" collectionCode="LACM" pageId="51" pageNumber="52" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
) that is almost certainly an underestimate (see above), and two specimens (
|
||
<materialsCitation box="[1045,1176,1717,1744]" collectionCode="MOR" pageId="51" pageNumber="52" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, TMP 1981.006.0001) that have the same LAG count of 18. Therefore, chronological age can be used as a proxy for relative maturity throughout the growth series as long as sampling from a skeleton is sufficiently thorough (cf.
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[661,926,1837,1863]" journalOrPublisher="Science Advances" pageId="51" pageNumber="52" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[661,926,1837,1863]" italics="true" pageId="51" pageNumber="52">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825004" ID-Zenodo-Dep="5825004" httpUri="https://zenodo.org/record/5825004/files/figure.png" pageId="52" pageNumber="53" startId="52.[524,589,704,726]" targetBox="[498,1541,234,691]" targetPageId="52">
|
||
<paragraph blockId="52.[524,1515,704,814]" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" pageId="52" pageNumber="53">
|
||
Figure 16 Bivariate scatterplot showing the relationship between chronological age with maturity among eight specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[805,995,734,755]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="52" pageNumber="53" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[805,995,734,755]" italics="true" pageId="52" pageNumber="53">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The comparison is limited to specimens that have been histologically aged; growth stages (
|
||
<emphasis box="[915,926,764,785]" italics="true" pageId="52" pageNumber="53">x</emphasis>
|
||
-axis) and chronological age (
|
||
<emphasis box="[1217,1227,764,785]" italics="true" pageId="52" pageNumber="53">y</emphasis>
|
||
-axis) have been converted to ranks. See Table 14 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-16
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFF0FF9CFDEBFCF73331FC73" httpUri="http://table.plazi.org/id/DF5C84E2FFF0FF9CFDEBFCF73331FC73" pageId="52" pageNumber="53" startId="52.[525,580,895,916]" targetBox="[516,1497,996,1351]" targetIsTable="true" targetPageId="52">
|
||
<paragraph blockId="52.[525,1515,895,975]" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[525,1359,895,917]" pageId="52" pageNumber="53">
|
||
Table 14 Summary of chronological age data and growth rank in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1168,1353,895,916]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="52" pageNumber="53" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1168,1353,895,916]" italics="true" pageId="52" pageNumber="53">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of the raw and ranked data used in the Spearman correlation test of maturity and chronological age in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1454,1510,925,946]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="52" pageNumber="53" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1454,1510,925,946]" italics="true" pageId="52" pageNumber="53">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="52" pageNumber="53">
|
||
<table box="[516,1497,996,1351]" gridcols="6" gridrows="9" pageId="52" pageNumber="53">
|
||
<tr box="[516,1497,996,1047]" gridrow="0" pageId="52" pageNumber="53">
|
||
<th box="[516,709,996,1047]" gridcol="0" gridrow="0" pageId="52" pageNumber="53">Specimen</th>
|
||
<th box="[725,867,996,1047]" gridcol="1" gridrow="0" pageId="52" pageNumber="53">Chronological age</th>
|
||
<th box="[890,1032,996,1047]" gridcol="2" gridrow="0" pageId="52" pageNumber="53">Chronological age rank</th>
|
||
<th box="[1070,1252,996,1047]" gridcol="3" gridrow="0" pageId="52" pageNumber="53">Chronological age midranks</th>
|
||
<th box="[1285,1361,996,1047]" gridcol="4" gridrow="0" pageId="52" pageNumber="53">Growth stage</th>
|
||
<th box="[1394,1497,996,1047]" gridcol="5" gridrow="0" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1497,996,1046]" pageId="52" pageNumber="53">Growth stage rank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1497,1066,1087]" gridrow="1" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1066,1087]" gridcol="0" gridrow="1" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,646,1066,1087]" collectionCode="LACM" pageId="52" pageNumber="53" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1066,1087]" gridcol="1" gridrow="1" pageId="52" pageNumber="53">2</td>
|
||
<td box="[890,1032,1066,1087]" gridcol="2" gridrow="1" pageId="52" pageNumber="53">1</td>
|
||
<td box="[1070,1252,1066,1087]" gridcol="3" gridrow="1" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1066,1087]" pageId="52" pageNumber="53">1</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1066,1087]" gridcol="4" gridrow="1" pageId="52" pageNumber="53">1</td>
|
||
<td box="[1394,1497,1066,1087]" gridcol="5" gridrow="1" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1066,1087]" pageId="52" pageNumber="53">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1104,1125]" gridrow="2" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1104,1125]" gridcol="0" gridrow="2" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,665,1104,1125]" collectionCode="BMRP" pageId="52" pageNumber="53" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1104,1125]" gridcol="1" gridrow="2" pageId="52" pageNumber="53">13</td>
|
||
<td box="[890,1032,1104,1125]" gridcol="2" gridrow="2" pageId="52" pageNumber="53">2</td>
|
||
<td box="[1070,1252,1104,1125]" gridcol="3" gridrow="2" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1104,1125]" pageId="52" pageNumber="53">2</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1104,1125]" gridcol="4" gridrow="2" pageId="52" pageNumber="53">4</td>
|
||
<td box="[1394,1497,1104,1125]" gridcol="5" gridrow="2" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1104,1125]" pageId="52" pageNumber="53">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1141,1163]" gridrow="3" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1141,1163]" gridcol="0" gridrow="3" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,665,1141,1163]" collectionCode="BMRP" pageId="52" pageNumber="53" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1141,1163]" gridcol="1" gridrow="3" pageId="52" pageNumber="53">15</td>
|
||
<td box="[890,1032,1141,1163]" gridcol="2" gridrow="3" pageId="52" pageNumber="53">4</td>
|
||
<td box="[1070,1252,1141,1163]" gridcol="3" gridrow="3" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1141,1162]" pageId="52" pageNumber="53">4</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1141,1163]" gridcol="4" gridrow="3" pageId="52" pageNumber="53">5</td>
|
||
<td box="[1394,1497,1141,1163]" gridcol="5" gridrow="3" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1141,1162]" pageId="52" pageNumber="53">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1179,1201]" gridrow="4" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1179,1201]" gridcol="0" gridrow="4" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,646,1179,1201]" collectionCode="LACM" pageId="52" pageNumber="53" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1179,1201]" gridcol="1" gridrow="4" pageId="52" pageNumber="53">14</td>
|
||
<td box="[890,1032,1179,1201]" gridcol="2" gridrow="4" pageId="52" pageNumber="53">3</td>
|
||
<td box="[1070,1252,1179,1201]" gridcol="3" gridrow="4" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1179,1200]" pageId="52" pageNumber="53">3</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1179,1201]" gridcol="4" gridrow="4" pageId="52" pageNumber="53">6</td>
|
||
<td box="[1394,1497,1179,1201]" gridcol="5" gridrow="4" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1179,1200]" pageId="52" pageNumber="53">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1216,1238]" gridrow="5" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1216,1238]" gridcol="0" gridrow="5" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,622,1216,1238]" collectionCode="MOR" pageId="52" pageNumber="53" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1216,1238]" gridcol="1" gridrow="5" pageId="52" pageNumber="53">18</td>
|
||
<td box="[890,1032,1216,1238]" gridcol="2" gridrow="5" pageId="52" pageNumber="53">5</td>
|
||
<td box="[1070,1252,1216,1238]" gridcol="3" gridrow="5" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1100,1217,1238]" pageId="52" pageNumber="53">5.5</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1216,1238]" gridcol="4" gridrow="5" pageId="52" pageNumber="53">7</td>
|
||
<td box="[1394,1497,1216,1238]" gridcol="5" gridrow="5" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1217,1238]" pageId="52" pageNumber="53">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1254,1276]" gridrow="6" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1254,1276]" gridcol="0" gridrow="6" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,709,1254,1276]" collectionCode="TMP" pageId="52" pageNumber="53" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1254,1276]" gridcol="1" gridrow="6" pageId="52" pageNumber="53">18</td>
|
||
<td box="[890,1032,1254,1276]" gridcol="2" gridrow="6" pageId="52" pageNumber="53">6</td>
|
||
<td box="[1070,1252,1254,1276]" gridcol="3" gridrow="6" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1100,1254,1275]" pageId="52" pageNumber="53">5.5</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1254,1276]" gridcol="4" gridrow="6" pageId="52" pageNumber="53">8</td>
|
||
<td box="[1394,1497,1254,1276]" gridcol="5" gridrow="6" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1254,1275]" pageId="52" pageNumber="53">6</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1292,1313]" gridrow="7" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1292,1313]" gridcol="0" gridrow="7" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,709,1292,1313]" collectionCode="TMP" pageId="52" pageNumber="53" specimenCode="TMP 1981.006.0001">TMP 1981.012.0001</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1292,1313]" gridcol="1" gridrow="7" pageId="52" pageNumber="53">22</td>
|
||
<td box="[890,1032,1292,1313]" gridcol="2" gridrow="7" pageId="52" pageNumber="53">7</td>
|
||
<td box="[1070,1252,1292,1313]" gridcol="3" gridrow="7" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1292,1313]" pageId="52" pageNumber="53">7</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1292,1313]" gridcol="4" gridrow="7" pageId="52" pageNumber="53">11</td>
|
||
<td box="[1394,1497,1292,1313]" gridcol="5" gridrow="7" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1292,1313]" pageId="52" pageNumber="53">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1497,1329,1351]" gridrow="8" pageId="52" pageNumber="53">
|
||
<th box="[516,709,1329,1351]" gridcol="0" gridrow="8" pageId="52" pageNumber="53">
|
||
<materialsCitation box="[516,667,1329,1351]" collectionCode="FMNH" pageId="52" pageNumber="53" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[725,867,1329,1351]" gridcol="1" gridrow="8" pageId="52" pageNumber="53">28</td>
|
||
<td box="[890,1032,1329,1351]" gridcol="2" gridrow="8" pageId="52" pageNumber="53">8</td>
|
||
<td box="[1070,1252,1329,1351]" gridcol="3" gridrow="8" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1070,1082,1329,1350]" pageId="52" pageNumber="53">8</emphasis>
|
||
</td>
|
||
<td box="[1285,1361,1329,1351]" gridcol="4" gridrow="8" pageId="52" pageNumber="53">20</td>
|
||
<td box="[1394,1497,1329,1351]" gridcol="5" gridrow="8" pageId="52" pageNumber="53">
|
||
<emphasis bold="true" box="[1394,1406,1329,1350]" pageId="52" pageNumber="53">8</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="52.[498,1542,1418,1927]" box="[498,782,1418,1446]" pageId="52" pageNumber="53">
|
||
<heading bold="true" box="[498,782,1418,1446]" fontSize="11" level="3" pageId="52" pageNumber="53" reason="6">
|
||
<emphasis bold="true" box="[498,782,1418,1446]" italics="true" pageId="52" pageNumber="53">Correlation with size</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="52.[498,1542,1418,1927]" lastBlockId="53.[498,1513,1678,1745]" lastPageId="53" lastPageNumber="54" pageId="52" pageNumber="53">
|
||
The scatterplot between maturity and size of ranked data shows an overall increasing trend, but with variation among the adult specimens (
|
||
<figureCitation box="[1133,1211,1501,1528]" captionStart="Figure 17" captionStartId="53.[524,589,712,734]" captionTargetBox="[504,1453,237,675]" captionTargetId="figure-84@53.[498,1458,234,685]" captionTargetPageId="53" captionText="Figure 17 Bivariate scatterplots showing the relationship between size with maturity among 15 specimens of Tyrannosaurus rex. The comparison is limited to specimens that have comparable size data; growth stages (x-axis) and size (y-axis) have been converted to ranks. See Table 15 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-17" figureDoi="http://doi.org/10.5281/zenodo.5825006" httpUri="https://zenodo.org/record/5825006/files/figure.png" pageId="52" pageNumber="53">Fig. 17</figureCitation>
|
||
;
|
||
<tableCitation box="[1226,1327,1501,1528]" captionStart="Table 15" captionStartId="53.[525,580,891,912]" captionTargetBox="[516,1525,992,1611]" captionText="Table 15 Summary of size data and growth rank in Tyrannosaurus rex. Summary of raw and ranked data used in the Spearman correlation test of size rank and growth rank in T. rex. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FFF1FF9DFDEBFCF3344BFC77" pageId="52" pageNumber="53" tableUuid="DF5C84E2FFF1FF9DFDEBFCF3344BFC77">Table 15</tableCitation>
|
||
). A Shapiro
|
||
<emphasis box="[1469,1484,1502,1527]" italics="true" pageId="52" pageNumber="53">–</emphasis>
|
||
Wilk test of normality found that the ranked size data and growth rank data are normally distributed (
|
||
<emphasis box="[641,655,1582,1607]" italics="true" pageId="52" pageNumber="53">p</emphasis>
|
||
= 0.166 and 0.678, respectively). The Spearman rank correlation test resulted in a significant (
|
||
<emphasis box="[691,705,1622,1647]" italics="true" pageId="52" pageNumber="53">p</emphasis>
|
||
= 0.000) correlation (
|
||
<emphasis box="[966,977,1622,1647]" italics="true" pageId="52" pageNumber="53">
|
||
<emphasis box="[977,987,1633,1649]" italics="true" pageId="52" pageNumber="53">
|
||
<subScript attach="left" box="[977,987,1633,1649]" fontSize="7" pageId="52" pageNumber="53">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.903), showing that size can serve as a proxy for maturity; however, this congruence is seen among animals that are less than adult size. A second test was run on the animals in the adult size range (growth ranks 5
|
||
<emphasis box="[512,527,1741,1766]" italics="true" pageId="52" pageNumber="53">–</emphasis>
|
||
15), where a significant (
|
||
<emphasis box="[811,825,1741,1766]" italics="true" pageId="52" pageNumber="53">p</emphasis>
|
||
= 0.008) correlation (
|
||
<emphasis box="[1078,1089,1741,1766]" italics="true" pageId="52" pageNumber="53">r</emphasis>
|
||
<subScript attach="left" box="[1089,1099,1753,1770]" fontSize="7" pageId="52" pageNumber="53">S</subScript>
|
||
= 0.748) was obtained, indicating that among adult-sized animals, size is a reliable proxy for maturity. Even with
|
||
<materialsCitation box="[1384,1516,1780,1807]" collectionCode="MOR" pageId="52" pageNumber="53" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, a relatively small young adult (femur length: 1.16 m), excluded (limiting the comparison to growth ranks 6
|
||
<emphasis box="[675,690,1861,1886]" italics="true" pageId="52" pageNumber="53">–</emphasis>
|
||
15), the results are significant (
|
||
<emphasis box="[1051,1062,1861,1886]" italics="true" pageId="52" pageNumber="53">r</emphasis>
|
||
<subScript attach="left" box="[1062,1072,1872,1889]" fontSize="7" pageId="52" pageNumber="53">S</subScript>
|
||
= 0.661,
|
||
<emphasis box="[1182,1196,1861,1886]" italics="true" pageId="52" pageNumber="53">p</emphasis>
|
||
= 0.038). However, when the test is limited to large adults (skull length greater than 1.2 m; growth ranks 8
|
||
<emphasis box="[1408,1423,1901,1926]" italics="true" pageId="52" pageNumber="53">–</emphasis>
|
||
15) the results are nonsignificant (
|
||
<emphasis box="[812,823,1679,1704]" italics="true" pageId="53" pageNumber="54">r</emphasis>
|
||
<subScript attach="left" box="[823,833,1691,1708]" fontSize="7" pageId="53" pageNumber="54">S</subScript>
|
||
= 0.312,
|
||
<emphasis box="[945,959,1679,1704]" italics="true" pageId="53" pageNumber="54">p</emphasis>
|
||
= 0.451), indicating that size cannot serve as a proxy for maturity among the largest adults.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825006" ID-Zenodo-Dep="5825006" httpUri="https://zenodo.org/record/5825006/files/figure.png" pageId="53" pageNumber="54" startId="53.[524,589,712,734]" targetBox="[504,1453,237,675]" targetPageId="53">
|
||
<paragraph blockId="53.[524,1516,712,822]" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" pageId="53" pageNumber="54">
|
||
Figure 17 Bivariate scatterplots showing the relationship between size with maturity among 15 specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[662,850,742,763]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="53" pageNumber="54" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[662,850,742,763]" italics="true" pageId="53" pageNumber="54">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The comparison is limited to specimens that have comparable size data; growth stages (
|
||
<emphasis box="[727,738,772,793]" italics="true" pageId="53" pageNumber="54">x</emphasis>
|
||
-axis) and size (
|
||
<emphasis box="[897,907,772,793]" italics="true" pageId="53" pageNumber="54">y</emphasis>
|
||
-axis) have been converted to ranks. See Table 15 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-17
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFF1FF9DFDEBFCF3344BFC77" httpUri="http://table.plazi.org/id/DF5C84E2FFF1FF9DFDEBFCF3344BFC77" pageId="53" pageNumber="54" startId="53.[525,580,891,912]" targetBox="[516,1525,992,1611]" targetIsTable="true" targetPageId="53">
|
||
<paragraph blockId="53.[525,1515,891,972]" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[525,1235,891,913]" pageId="53" pageNumber="54">
|
||
Table 15 Summary of size data and growth rank in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1043,1229,891,912]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="53" pageNumber="54" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1043,1229,891,912]" italics="true" pageId="53" pageNumber="54">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of raw and ranked data used in the Spearman correlation test of size rank and growth rank in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1241,1295,921,942]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="53" pageNumber="54" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1241,1295,921,942]" italics="true" pageId="53" pageNumber="54">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="53" pageNumber="54">
|
||
<table box="[516,1525,992,1611]" gridcols="7" gridrows="16" pageId="53" pageNumber="54">
|
||
<tr box="[516,1525,992,1043]" gridrow="0" pageId="53" pageNumber="54">
|
||
<th box="[516,709,992,1043]" gridcol="0" gridrow="0" pageId="53" pageNumber="54">Specimen</th>
|
||
<th box="[725,771,992,1043]" gridcol="1" gridrow="0" pageId="53" pageNumber="54">Size (cm)</th>
|
||
<th box="[825,871,992,1043]" gridcol="2" gridrow="0" pageId="53" pageNumber="54">Size rank</th>
|
||
<th box="[925,1020,992,1043]" gridcol="3" gridrow="0" pageId="53" pageNumber="54">Size midranks</th>
|
||
<th box="[1070,1146,992,1043]" gridcol="4" gridrow="0" pageId="53" pageNumber="54">Growth stage</th>
|
||
<th box="[1207,1340,992,1043]" gridcol="5" gridrow="0" pageId="53" pageNumber="54">Growth stage rank</th>
|
||
<th box="[1392,1525,992,1043]" gridcol="6" gridrow="0" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" pageId="53" pageNumber="54">Growth stage midranks</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1525,1062,1084]" gridrow="1" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1062,1084]" gridcol="0" gridrow="1" pageId="53" pageNumber="54">
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<materialsCitation box="[516,646,1063,1084]" collectionCode="LACM" pageId="53" pageNumber="54" specimenCode="LACM 28471">LACM 28471</materialsCitation>
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||
</th>
|
||
<td box="[725,771,1062,1084]" gridcol="1" gridrow="1" pageId="53" pageNumber="54">40</td>
|
||
<td box="[825,871,1062,1084]" gridcol="2" gridrow="1" pageId="53" pageNumber="54">1</td>
|
||
<td box="[925,1020,1062,1084]" gridcol="3" gridrow="1" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,937,1062,1083]" pageId="53" pageNumber="54">1</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1062,1084]" gridcol="4" gridrow="1" pageId="53" pageNumber="54">1</td>
|
||
<td box="[1207,1340,1062,1084]" gridcol="5" gridrow="1" pageId="53" pageNumber="54">1</td>
|
||
<td box="[1392,1525,1062,1084]" gridcol="6" gridrow="1" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1062,1083]" pageId="53" pageNumber="54">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1100,1122]" gridrow="2" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1100,1122]" gridcol="0" gridrow="2" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,642,1100,1122]" collectionCode="CMNH" pageId="53" pageNumber="54" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
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||
</th>
|
||
<td box="[725,771,1100,1122]" gridcol="1" gridrow="2" pageId="53" pageNumber="54">57</td>
|
||
<td box="[825,871,1100,1122]" gridcol="2" gridrow="2" pageId="53" pageNumber="54">2</td>
|
||
<td box="[925,1020,1100,1122]" gridcol="3" gridrow="2" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,937,1100,1121]" pageId="53" pageNumber="54">2</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1100,1122]" gridcol="4" gridrow="2" pageId="53" pageNumber="54">3</td>
|
||
<td box="[1207,1340,1100,1122]" gridcol="5" gridrow="2" pageId="53" pageNumber="54">2</td>
|
||
<td box="[1392,1525,1100,1122]" gridcol="6" gridrow="2" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1100,1121]" pageId="53" pageNumber="54">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1137,1159]" gridrow="3" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1137,1159]" gridcol="0" gridrow="3" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,665,1138,1159]" collectionCode="BMRP" pageId="53" pageNumber="54" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1137,1159]" gridcol="1" gridrow="3" pageId="53" pageNumber="54">74</td>
|
||
<td box="[825,871,1137,1159]" gridcol="2" gridrow="3" pageId="53" pageNumber="54">3</td>
|
||
<td box="[925,1020,1137,1159]" gridcol="3" gridrow="3" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,937,1137,1158]" pageId="53" pageNumber="54">3</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1137,1159]" gridcol="4" gridrow="3" pageId="53" pageNumber="54">4</td>
|
||
<td box="[1207,1340,1137,1159]" gridcol="5" gridrow="3" pageId="53" pageNumber="54">3</td>
|
||
<td box="[1392,1525,1137,1159]" gridcol="6" gridrow="3" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1137,1158]" pageId="53" pageNumber="54">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1175,1197]" gridrow="4" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1175,1197]" gridcol="0" gridrow="4" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,646,1175,1197]" collectionCode="LACM" pageId="53" pageNumber="54" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1175,1197]" gridcol="1" gridrow="4" pageId="53" pageNumber="54">80</td>
|
||
<td box="[825,871,1175,1197]" gridcol="2" gridrow="4" pageId="53" pageNumber="54">4</td>
|
||
<td box="[925,1020,1175,1197]" gridcol="3" gridrow="4" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,937,1175,1196]" pageId="53" pageNumber="54">4</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1175,1197]" gridcol="4" gridrow="4" pageId="53" pageNumber="54">6</td>
|
||
<td box="[1207,1340,1175,1197]" gridcol="5" gridrow="4" pageId="53" pageNumber="54">4</td>
|
||
<td box="[1392,1525,1175,1197]" gridcol="6" gridrow="4" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1175,1196]" pageId="53" pageNumber="54">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1213,1235]" gridrow="5" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1213,1235]" gridcol="0" gridrow="5" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,622,1213,1235]" collectionCode="MOR" pageId="53" pageNumber="54" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1213,1235]" gridcol="1" gridrow="5" pageId="53" pageNumber="54">116</td>
|
||
<td box="[825,871,1213,1235]" gridcol="2" gridrow="5" pageId="53" pageNumber="54">5</td>
|
||
<td box="[925,1020,1213,1235]" gridcol="3" gridrow="5" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,937,1213,1234]" pageId="53" pageNumber="54">5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1213,1235]" gridcol="4" gridrow="5" pageId="53" pageNumber="54">7</td>
|
||
<td box="[1207,1340,1213,1235]" gridcol="5" gridrow="5" pageId="53" pageNumber="54">5</td>
|
||
<td box="[1392,1525,1213,1235]" gridcol="6" gridrow="5" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1213,1234]" pageId="53" pageNumber="54">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1250,1272]" gridrow="6" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1250,1272]" gridcol="0" gridrow="6" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,709,1250,1272]" collectionCode="TMP" pageId="53" pageNumber="54" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1250,1272]" gridcol="1" gridrow="6" pageId="53" pageNumber="54">120</td>
|
||
<td box="[825,871,1250,1272]" gridcol="2" gridrow="6" pageId="53" pageNumber="54">6</td>
|
||
<td box="[925,1020,1250,1272]" gridcol="3" gridrow="6" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,955,1250,1271]" pageId="53" pageNumber="54">6.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1250,1272]" gridcol="4" gridrow="6" pageId="53" pageNumber="54">8</td>
|
||
<td box="[1207,1340,1250,1272]" gridcol="5" gridrow="6" pageId="53" pageNumber="54">6</td>
|
||
<td box="[1392,1525,1250,1272]" gridcol="6" gridrow="6" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1250,1271]" pageId="53" pageNumber="54">6</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1288,1310]" gridrow="7" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1288,1310]" gridcol="0" gridrow="7" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,658,1288,1310]" collectionCode="LACM" pageId="53" pageNumber="54" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1288,1310]" gridcol="1" gridrow="7" pageId="53" pageNumber="54">120</td>
|
||
<td box="[825,871,1288,1310]" gridcol="2" gridrow="7" pageId="53" pageNumber="54">7</td>
|
||
<td box="[925,1020,1288,1310]" gridcol="3" gridrow="7" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,955,1288,1309]" pageId="53" pageNumber="54">6.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1288,1310]" gridcol="4" gridrow="7" pageId="53" pageNumber="54">9</td>
|
||
<td box="[1207,1340,1288,1310]" gridcol="5" gridrow="7" pageId="53" pageNumber="54">7</td>
|
||
<td box="[1392,1525,1288,1310]" gridcol="6" gridrow="7" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1288,1309]" pageId="53" pageNumber="54">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1326,1347]" gridrow="8" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1326,1347]" gridcol="0" gridrow="8" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,656,1326,1347]" collectionCode="UWBM" pageId="53" pageNumber="54" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1326,1347]" gridcol="1" gridrow="8" pageId="53" pageNumber="54">130</td>
|
||
<td box="[825,871,1326,1347]" gridcol="2" gridrow="8" pageId="53" pageNumber="54">8</td>
|
||
<td box="[925,1020,1326,1347]" gridcol="3" gridrow="8" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,955,1326,1347]" pageId="53" pageNumber="54">8.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1326,1347]" gridcol="4" gridrow="8" pageId="53" pageNumber="54">11</td>
|
||
<td box="[1207,1340,1326,1347]" gridcol="5" gridrow="8" pageId="53" pageNumber="54">8</td>
|
||
<td box="[1392,1525,1326,1347]" gridcol="6" gridrow="8" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1326,1347]" pageId="53" pageNumber="54">8</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1363,1385]" gridrow="9" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1363,1385]" gridcol="0" gridrow="9" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,632,1363,1385]" collectionCode="RSM" pageId="53" pageNumber="54" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1363,1385]" gridcol="1" gridrow="9" pageId="53" pageNumber="54">140</td>
|
||
<td box="[825,871,1363,1385]" gridcol="2" gridrow="9" pageId="53" pageNumber="54">12</td>
|
||
<td box="[925,1020,1363,1385]" gridcol="3" gridrow="9" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1363,1384]" pageId="53" pageNumber="54">13.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1363,1385]" gridcol="4" gridrow="9" pageId="53" pageNumber="54">12</td>
|
||
<td box="[1207,1340,1363,1385]" gridcol="5" gridrow="9" pageId="53" pageNumber="54">9</td>
|
||
<td box="[1392,1525,1363,1385]" gridcol="6" gridrow="9" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1404,1363,1384]" pageId="53" pageNumber="54">9</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1401,1423]" gridrow="10" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1401,1423]" gridcol="0" gridrow="10" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,707,1401,1423]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="53" pageNumber="54" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1401,1423]" gridcol="1" gridrow="10" pageId="53" pageNumber="54">136</td>
|
||
<td box="[825,871,1401,1423]" gridcol="2" gridrow="10" pageId="53" pageNumber="54">10</td>
|
||
<td box="[925,1020,1401,1423]" gridcol="3" gridrow="10" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1401,1422]" pageId="53" pageNumber="54">10.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1401,1423]" gridcol="4" gridrow="10" pageId="53" pageNumber="54">13</td>
|
||
<td box="[1207,1340,1401,1423]" gridcol="5" gridrow="10" pageId="53" pageNumber="54">10</td>
|
||
<td box="[1392,1525,1401,1423]" gridcol="6" gridrow="10" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1416,1401,1422]" pageId="53" pageNumber="54">10</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1438,1460]" gridrow="11" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1438,1460]" gridcol="0" gridrow="11" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,610,1439,1460]" collectionCode="MOR" pageId="53" pageNumber="54" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1438,1460]" gridcol="1" gridrow="11" pageId="53" pageNumber="54">136</td>
|
||
<td box="[825,871,1438,1460]" gridcol="2" gridrow="11" pageId="53" pageNumber="54">11</td>
|
||
<td box="[925,1020,1438,1460]" gridcol="3" gridrow="11" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1438,1459]" pageId="53" pageNumber="54">10.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1438,1460]" gridcol="4" gridrow="11" pageId="53" pageNumber="54">16</td>
|
||
<td box="[1207,1340,1438,1460]" gridcol="5" gridrow="11" pageId="53" pageNumber="54">11</td>
|
||
<td box="[1392,1525,1438,1460]" gridcol="6" gridrow="11" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1434,1438,1459]" pageId="53" pageNumber="54">11.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1476,1498]" gridrow="12" pageId="53" pageNumber="54">
|
||
<th box="[516,709,1476,1498]" gridcol="0" gridrow="12" pageId="53" pageNumber="54">
|
||
<materialsCitation box="[516,646,1476,1497]" collectionCode="LACM" pageId="53" pageNumber="54" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[725,771,1476,1498]" gridcol="1" gridrow="12" pageId="53" pageNumber="54">140</td>
|
||
<td box="[825,871,1476,1498]" gridcol="2" gridrow="12" pageId="53" pageNumber="54">14</td>
|
||
<td box="[925,1020,1476,1498]" gridcol="3" gridrow="12" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1476,1497]" pageId="53" pageNumber="54">13.5</emphasis>
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||
</td>
|
||
<td box="[1070,1146,1476,1498]" gridcol="4" gridrow="12" pageId="53" pageNumber="54">16</td>
|
||
<td box="[1207,1340,1476,1498]" gridcol="5" gridrow="12" pageId="53" pageNumber="54">12</td>
|
||
<td box="[1392,1525,1476,1498]" gridcol="6" gridrow="12" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1434,1476,1497]" pageId="53" pageNumber="54">11.5</emphasis>
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||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1513,1535]" gridrow="13" pageId="53" pageNumber="54">
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<th box="[516,709,1513,1535]" gridcol="0" gridrow="13" pageId="53" pageNumber="54">
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<materialsCitation box="[516,606,1514,1535]" collectionCode="CM" pageId="53" pageNumber="54" specimenCode="CM 9380">CM 9380</materialsCitation>
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</th>
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<td box="[725,771,1513,1535]" gridcol="1" gridrow="13" pageId="53" pageNumber="54">130</td>
|
||
<td box="[825,871,1513,1535]" gridcol="2" gridrow="13" pageId="53" pageNumber="54">9</td>
|
||
<td box="[925,1020,1513,1535]" gridcol="3" gridrow="13" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,955,1513,1534]" pageId="53" pageNumber="54">8.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1513,1535]" gridcol="4" gridrow="13" pageId="53" pageNumber="54">17</td>
|
||
<td box="[1207,1340,1513,1535]" gridcol="5" gridrow="13" pageId="53" pageNumber="54">13</td>
|
||
<td box="[1392,1525,1513,1535]" gridcol="6" gridrow="13" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1434,1513,1534]" pageId="53" pageNumber="54">13.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1551,1572]" gridrow="14" pageId="53" pageNumber="54">
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<th box="[516,709,1551,1572]" gridcol="0" gridrow="14" pageId="53" pageNumber="54">
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<materialsCitation box="[516,610,1551,1572]" collectionCode="MOR" pageId="53" pageNumber="54" specimenCode="MOR 008">MOR 008</materialsCitation>
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</th>
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<td box="[725,771,1551,1572]" gridcol="1" gridrow="14" pageId="53" pageNumber="54">140</td>
|
||
<td box="[825,871,1551,1572]" gridcol="2" gridrow="14" pageId="53" pageNumber="54">13</td>
|
||
<td box="[925,1020,1551,1572]" gridcol="3" gridrow="14" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1551,1572]" pageId="53" pageNumber="54">13.5</emphasis>
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||
</td>
|
||
<td box="[1070,1146,1551,1572]" gridcol="4" gridrow="14" pageId="53" pageNumber="54">17</td>
|
||
<td box="[1207,1340,1551,1572]" gridcol="5" gridrow="14" pageId="53" pageNumber="54">14</td>
|
||
<td box="[1392,1525,1551,1572]" gridcol="6" gridrow="14" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1434,1551,1572]" pageId="53" pageNumber="54">13.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1589,1611]" gridrow="15" pageId="53" pageNumber="54">
|
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<th box="[516,709,1589,1611]" gridcol="0" gridrow="15" pageId="53" pageNumber="54">
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<materialsCitation box="[516,667,1589,1610]" collectionCode="FMNH" pageId="53" pageNumber="54" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
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</th>
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<td box="[725,771,1589,1611]" gridcol="1" gridrow="15" pageId="53" pageNumber="54">140</td>
|
||
<td box="[825,871,1589,1611]" gridcol="2" gridrow="15" pageId="53" pageNumber="54">15</td>
|
||
<td box="[925,1020,1589,1611]" gridcol="3" gridrow="15" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[925,967,1589,1610]" pageId="53" pageNumber="54">13.5</emphasis>
|
||
</td>
|
||
<td box="[1070,1146,1589,1611]" gridcol="4" gridrow="15" pageId="53" pageNumber="54">20</td>
|
||
<td box="[1207,1340,1589,1611]" gridcol="5" gridrow="15" pageId="53" pageNumber="54">15</td>
|
||
<td box="[1392,1525,1589,1611]" gridcol="6" gridrow="15" pageId="53" pageNumber="54">
|
||
<emphasis bold="true" box="[1392,1416,1589,1610]" pageId="53" pageNumber="54">15</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
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</paragraph>
|
||
<paragraph blockId="53.[498,1542,1783,1934]" box="[498,801,1783,1811]" pageId="53" pageNumber="54">
|
||
<heading bold="true" box="[498,801,1783,1811]" fontSize="11" level="3" pageId="53" pageNumber="54" reason="6">
|
||
<emphasis bold="true" box="[498,801,1783,1811]" italics="true" pageId="53" pageNumber="54">Correlation with mass</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="53.[498,1542,1783,1934]" lastBlockId="54.[498,1542,1507,1732]" lastPageId="54" lastPageNumber="55" pageId="53" pageNumber="54">
|
||
The scatterplot between congruence between maturity and mass of ranked data (
|
||
<figureCitation box="[1453,1533,1827,1853]" captionStart="Figure 18" captionStartId="54.[524,589,709,731]" captionTargetBox="[506,1455,237,675]" captionTargetId="figure-144@54.[498,1458,234,682]" captionTargetPageId="54" captionText="Figure 18 Bivariate scatterplots showing the relationship between mass with maturity among nine specimens of Tyrannosaurus rex. The comparison is limited to specimens that have published mass estimates; growth stages (x-axis) and mass (y-axis) have been converted to ranks. See Table 16 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-18" figureDoi="http://doi.org/10.5281/zenodo.5825008" httpUri="https://zenodo.org/record/5825008/files/figure.png" pageId="53" pageNumber="54">Fig. 18</figureCitation>
|
||
;
|
||
<tableCitation box="[498,600,1867,1893]" captionStart="Table 16" captionStartId="54.[525,580,909,930]" captionTargetBox="[516,1525,1038,1432]" captionText="Table 16 Summary of mass data and growth rank for Tyrannosaurus rex. Summary of raw and ranked data for the Spearman correlation test of mass and maturity in T. rex. Masses are drawn from Erickson et al. (2004) and Persons, Currie & Erickson (2019). Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FFF2FF9EFDEBFC05352EFC46" pageId="53" pageNumber="54" tableUuid="DF5C84E2FFF2FF9EFDEBFC05352EFC46">Table 16</tableCitation>
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||
) shows a cloud of points with an increasing trend, indicating correlation. A Shapiro
|
||
<emphasis box="[616,631,1908,1933]" italics="true" pageId="53" pageNumber="54">–</emphasis>
|
||
Wilk test found that the ranked mass data and growth rank data are normally distributed (
|
||
<emphasis box="[641,655,1508,1533]" italics="true" pageId="54" pageNumber="55">p</emphasis>
|
||
= 0.914 and 0.870, respectively). The Spearman correlation rank test resulted in a significant (
|
||
<emphasis box="[691,705,1548,1573]" italics="true" pageId="54" pageNumber="55">p</emphasis>
|
||
= 0.019) correlation (
|
||
<emphasis box="[966,977,1548,1573]" italics="true" pageId="54" pageNumber="55">
|
||
<emphasis box="[977,987,1559,1575]" italics="true" pageId="54" pageNumber="55">
|
||
<subScript attach="left" box="[977,987,1559,1575]" fontSize="7" pageId="54" pageNumber="55">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.753) between the variables; a second run of the test was made for the adult-sized animals (growth stage ranks 3
|
||
<emphasis box="[1307,1322,1588,1613]" italics="true" pageId="54" pageNumber="55">–</emphasis>
|
||
9) since this region of the distribution shows variation. The test resulted in a nonsignificant (
|
||
<emphasis box="[1364,1378,1627,1652]" italics="true" pageId="54" pageNumber="55">p</emphasis>
|
||
= 0.289) correlation (
|
||
<emphasis box="[644,655,1667,1692]" italics="true" pageId="54" pageNumber="55">
|
||
<emphasis box="[655,665,1679,1695]" italics="true" pageId="54" pageNumber="55">
|
||
<subScript attach="left" box="[655,665,1679,1695]" fontSize="7" pageId="54" pageNumber="55">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.468), indicating that mass is a poor predictor of maturity among the adult growth categories.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825008" ID-Zenodo-Dep="5825008" httpUri="https://zenodo.org/record/5825008/files/figure.png" pageId="54" pageNumber="55" startId="54.[524,589,709,731]" targetBox="[506,1455,237,675]" targetPageId="54">
|
||
<paragraph blockId="54.[524,1515,709,819]" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" pageId="54" pageNumber="55">
|
||
Figure 18 Bivariate scatterplots showing the relationship between mass with maturity among nine specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[663,851,739,760]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="55" phylum="Chordata" rank="species" species="rex">
|
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<emphasis bold="true" box="[663,851,739,760]" italics="true" pageId="54" pageNumber="55">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The comparison is limited to specimens that have published mass estimates; growth stages (
|
||
<emphasis box="[764,775,769,790]" italics="true" pageId="54" pageNumber="55">x</emphasis>
|
||
-axis) and mass (
|
||
<emphasis box="[935,945,769,790]" italics="true" pageId="54" pageNumber="55">y</emphasis>
|
||
-axis) have been converted to ranks. See Table 16 for the raw data. Full-size DOI: 10.7717/peerj.9192/fig-18
|
||
</paragraph>
|
||
</caption>
|
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<caption ID-Table-UUID="DF5C84E2FFF2FF9EFDEBFC05352EFC46" httpUri="http://table.plazi.org/id/DF5C84E2FFF2FF9EFDEBFC05352EFC46" pageId="54" pageNumber="55" startId="54.[525,580,909,930]" targetBox="[516,1525,1038,1432]" targetIsTable="true" targetPageId="54">
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<paragraph blockId="54.[525,1515,908,1018]" pageId="54" pageNumber="55">
|
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<emphasis bold="true" box="[525,1242,908,930]" pageId="54" pageNumber="55">
|
||
Table 16 Summary of mass data and growth rank for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1052,1236,909,930]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="55" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1052,1236,909,930]" italics="true" pageId="54" pageNumber="55">Tyrannosaurus rex</emphasis>
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||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Summary of raw and ranked data for the Spearman correlation test of mass and maturity in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1135,1190,938,959]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="54" pageNumber="55" phylum="Chordata" rank="species" species="rex">
|
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<emphasis box="[1135,1190,938,959]" italics="true" pageId="54" pageNumber="55">T. rex</emphasis>
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||
</taxonomicName>
|
||
. Masses are drawn from
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" journalOrPublisher="Nature" pageId="54" pageNumber="55" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
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<emphasis italics="true" pageId="54" pageNumber="55">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[688,1012,967,989]" journalOrPublisher="Anatomical Record" pageId="54" pageNumber="55" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
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<emphasis box="[688,1012,967,989]" italics="true" pageId="54" pageNumber="55">Persons, Currie & Erickson (2019)</emphasis>
|
||
</bibRefCitation>
|
||
. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="54" pageNumber="55">
|
||
<table box="[516,1525,1038,1432]" gridcols="6" gridrows="10" pageId="54" pageNumber="55">
|
||
<tr box="[516,1525,1038,1090]" gridrow="0" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1038,1090]" gridcol="0" gridrow="0" pageId="54" pageNumber="55">Specimen</th>
|
||
<th box="[761,858,1038,1090]" gridcol="1" gridrow="0" pageId="54" pageNumber="55">Mass (kg)</th>
|
||
<th box="[912,1014,1038,1090]" gridcol="2" gridrow="0" pageId="54" pageNumber="55">Mass rank</th>
|
||
<th box="[1069,1145,1038,1090]" gridcol="3" gridrow="0" pageId="54" pageNumber="55">Growth stage</th>
|
||
<th box="[1206,1338,1038,1090]" gridcol="4" gridrow="0" pageId="54" pageNumber="55">Growth stage rank</th>
|
||
<th box="[1393,1525,1038,1090]" gridcol="5" gridrow="0" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" pageId="54" pageNumber="55">Growth stage midranks</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1525,1109,1131]" gridrow="1" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1109,1131]" gridcol="0" gridrow="1" pageId="54" pageNumber="55">
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<materialsCitation box="[516,646,1109,1130]" collectionCode="LACM" pageId="54" pageNumber="55" specimenCode="LACM 28471">LACM 28471</materialsCitation>
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</th>
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<td box="[761,858,1109,1131]" gridcol="1" gridrow="1" pageId="54" pageNumber="55">25</td>
|
||
<td box="[912,1014,1109,1131]" gridcol="2" gridrow="1" pageId="54" pageNumber="55">1</td>
|
||
<td box="[1069,1145,1109,1131]" gridcol="3" gridrow="1" pageId="54" pageNumber="55">1</td>
|
||
<td box="[1206,1338,1109,1131]" gridcol="4" gridrow="1" pageId="54" pageNumber="55">1</td>
|
||
<td box="[1393,1525,1109,1131]" gridcol="5" gridrow="1" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1109,1130]" pageId="54" pageNumber="55">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1147,1169]" gridrow="2" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1147,1169]" gridcol="0" gridrow="2" pageId="54" pageNumber="55">
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<materialsCitation box="[516,646,1147,1169]" collectionCode="LACM" pageId="54" pageNumber="55" specimenCode="LACM 23845">LACM 23845</materialsCitation>
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</th>
|
||
<td box="[761,858,1147,1169]" gridcol="1" gridrow="2" pageId="54" pageNumber="55">98.9</td>
|
||
<td box="[912,1014,1147,1169]" gridcol="2" gridrow="2" pageId="54" pageNumber="55">2</td>
|
||
<td box="[1069,1145,1147,1169]" gridcol="3" gridrow="2" pageId="54" pageNumber="55">6</td>
|
||
<td box="[1206,1338,1147,1169]" gridcol="4" gridrow="2" pageId="54" pageNumber="55">2</td>
|
||
<td box="[1393,1525,1147,1169]" gridcol="5" gridrow="2" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1147,1168]" pageId="54" pageNumber="55">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1184,1206]" gridrow="3" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1184,1206]" gridcol="0" gridrow="3" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,622,1184,1206]" collectionCode="MOR" pageId="54" pageNumber="55" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[761,858,1184,1206]" gridcol="1" gridrow="3" pageId="54" pageNumber="55">6,100</td>
|
||
<td box="[912,1014,1184,1206]" gridcol="2" gridrow="3" pageId="54" pageNumber="55">5</td>
|
||
<td box="[1069,1145,1184,1206]" gridcol="3" gridrow="3" pageId="54" pageNumber="55">7</td>
|
||
<td box="[1206,1338,1184,1206]" gridcol="4" gridrow="3" pageId="54" pageNumber="55">3</td>
|
||
<td box="[1393,1525,1184,1206]" gridcol="5" gridrow="3" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1184,1205]" pageId="54" pageNumber="55">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1222,1244]" gridrow="4" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1222,1244]" gridcol="0" gridrow="4" pageId="54" pageNumber="55">
|
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<materialsCitation box="[516,707,1222,1244]" collectionCode="TMP" pageId="54" pageNumber="55" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[761,858,1222,1244]" gridcol="1" gridrow="4" pageId="54" pageNumber="55">4,469</td>
|
||
<td box="[912,1014,1222,1244]" gridcol="2" gridrow="4" pageId="54" pageNumber="55">3</td>
|
||
<td box="[1069,1145,1222,1244]" gridcol="3" gridrow="4" pageId="54" pageNumber="55">8</td>
|
||
<td box="[1206,1338,1222,1244]" gridcol="4" gridrow="4" pageId="54" pageNumber="55">4</td>
|
||
<td box="[1393,1525,1222,1244]" gridcol="5" gridrow="4" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1222,1243]" pageId="54" pageNumber="55">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1259,1281]" gridrow="5" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1259,1281]" gridcol="0" gridrow="5" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,632,1259,1281]" collectionCode="RSM" pageId="54" pageNumber="55" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[761,858,1259,1281]" gridcol="1" gridrow="5" pageId="54" pageNumber="55">8,870</td>
|
||
<td box="[912,1014,1259,1281]" gridcol="2" gridrow="5" pageId="54" pageNumber="55">9</td>
|
||
<td box="[1069,1145,1259,1281]" gridcol="3" gridrow="5" pageId="54" pageNumber="55">12</td>
|
||
<td box="[1206,1338,1259,1281]" gridcol="4" gridrow="5" pageId="54" pageNumber="55">5</td>
|
||
<td box="[1393,1525,1259,1281]" gridcol="5" gridrow="5" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1260,1281]" pageId="54" pageNumber="55">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1297,1319]" gridrow="6" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1297,1319]" gridcol="0" gridrow="6" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,610,1297,1319]" collectionCode="MOR" pageId="54" pageNumber="55" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[761,858,1297,1319]" gridcol="1" gridrow="6" pageId="54" pageNumber="55">6,264</td>
|
||
<td box="[912,1014,1297,1319]" gridcol="2" gridrow="6" pageId="54" pageNumber="55">6</td>
|
||
<td box="[1069,1145,1297,1319]" gridcol="3" gridrow="6" pageId="54" pageNumber="55">16</td>
|
||
<td box="[1206,1338,1297,1319]" gridcol="4" gridrow="6" pageId="54" pageNumber="55">6</td>
|
||
<td box="[1393,1525,1297,1319]" gridcol="5" gridrow="6" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1423,1297,1318]" pageId="54" pageNumber="55">6.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1334,1357]" gridrow="7" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1334,1357]" gridcol="0" gridrow="7" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,610,1335,1356]" collectionCode="MOR" pageId="54" pageNumber="55" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[761,858,1334,1357]" gridcol="1" gridrow="7" pageId="54" pageNumber="55">5,112</td>
|
||
<td box="[912,1014,1334,1357]" gridcol="2" gridrow="7" pageId="54" pageNumber="55">4</td>
|
||
<td box="[1069,1145,1334,1357]" gridcol="3" gridrow="7" pageId="54" pageNumber="55">16</td>
|
||
<td box="[1206,1338,1334,1357]" gridcol="4" gridrow="7" pageId="54" pageNumber="55">7</td>
|
||
<td box="[1393,1525,1334,1357]" gridcol="5" gridrow="7" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1423,1334,1355]" pageId="54" pageNumber="55">6.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1372,1394]" gridrow="8" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1372,1394]" gridcol="0" gridrow="8" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,606,1372,1393]" collectionCode="CM" pageId="54" pageNumber="55" specimenCode="CM 9380">CM 9380</materialsCitation>
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||
</th>
|
||
<td box="[761,858,1372,1394]" gridcol="1" gridrow="8" pageId="54" pageNumber="55">6,740</td>
|
||
<td box="[912,1014,1372,1394]" gridcol="2" gridrow="8" pageId="54" pageNumber="55">7</td>
|
||
<td box="[1069,1145,1372,1394]" gridcol="3" gridrow="8" pageId="54" pageNumber="55">17</td>
|
||
<td box="[1206,1338,1372,1394]" gridcol="4" gridrow="8" pageId="54" pageNumber="55">8</td>
|
||
<td box="[1393,1525,1372,1394]" gridcol="5" gridrow="8" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1372,1393]" pageId="54" pageNumber="55">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1410,1432]" gridrow="9" pageId="54" pageNumber="55">
|
||
<th box="[516,707,1410,1432]" gridcol="0" gridrow="9" pageId="54" pageNumber="55">
|
||
<materialsCitation box="[516,667,1410,1431]" collectionCode="FMNH" pageId="54" pageNumber="55" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
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||
</th>
|
||
<td box="[761,858,1410,1432]" gridcol="1" gridrow="9" pageId="54" pageNumber="55">8,462</td>
|
||
<td box="[912,1014,1410,1432]" gridcol="2" gridrow="9" pageId="54" pageNumber="55">8</td>
|
||
<td box="[1069,1145,1410,1432]" gridcol="3" gridrow="9" pageId="54" pageNumber="55">20</td>
|
||
<td box="[1206,1338,1410,1432]" gridcol="4" gridrow="9" pageId="54" pageNumber="55">9</td>
|
||
<td box="[1393,1525,1410,1432]" gridcol="5" gridrow="9" pageId="54" pageNumber="55">
|
||
<emphasis bold="true" box="[1393,1405,1410,1431]" pageId="54" pageNumber="55">8</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="54.[498,1541,1777,1929]" box="[498,1346,1777,1807]" pageId="54" pageNumber="55">
|
||
<heading bold="true" box="[498,1346,1777,1807]" fontSize="12" level="2" pageId="54" pageNumber="55" reason="0">
|
||
<emphasis bold="true" box="[498,1346,1777,1807]" pageId="54" pageNumber="55">Congruence between nonbiological factors and maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="54.[498,1541,1777,1929]" lastBlockId="56.[498,1539,1554,1700]" lastPageId="56" lastPageNumber="57" pageId="54" pageNumber="55">
|
||
The possibility that the growth series obtained here results from the influence of abiotic factors, namely geographic location and stratigraphic position, was tested using the Spearman rank correlation test since horizontal and vertical spatial data are hierarchical in time and space. For example, the growth series might reflect the north to south distribution of a latitudinally variable population, the anagenetic transformation of a population, or both trends. To test these hypotheses, geographic location was ranked by latitude along a north
|
||
<emphasis box="[564,579,1742,1767]" italics="true" pageId="55" pageNumber="56">–</emphasis>
|
||
south gradient that follows the long axis of Laramidia (
|
||
<tableCitation box="[1231,1333,1741,1767]" captionStart="Table 17" captionStartId="55.[525,580,248,269]" captionTargetBox="[516,1525,407,1523]" captionText="Table 17 Summary of data used in the correlation test between geographic latitude and maturity of Tyrannosaurus rex specimens. For each fossil, the columns are organized by specimen number, growth stage rank, geographic location rank, and level of resolution of the locality data. Specimens are listed by descending location rank (i.e., from north to south). Co, county; km, kilometer; m, meter; mi, miles; S, section; asterisk (*) indicates the resolution is almost certainly an underestimate." httpUri="http://table.plazi.org/id/DF5C84E2FFF3FF9FFDEBFF70320DFE3F" pageId="55" pageNumber="56" tableUuid="DF5C84E2FFF3FF9FFDEBFF70320DFE3F">Table 17</tableCitation>
|
||
;
|
||
<figureCitation box="[1348,1427,1741,1767]" captionStart="Figure 19" captionStartId="56.[524,589,731,753]" captionTargetBox="[497,1521,212,706]" captionTargetId="figure-133@56.[498,1458,234,704]" captionTargetPageId="56" captionText="Figure 19 Bivariate scatterplot showing the relationship between geographic location with maturity among 28 specimens Tyrannosaurus rex. Growth stages (x-axis) and geographic location (y-axis) have been converted to ranks. See Table 17 for the ranked data. “Montana North” refers to the region of Dawson, Garfield, and McCone counties, and “Montana South” refers to the region of Yellowstone and Carter counties. Maturity increases to the right along the x-axis; the y-axis follows the north-south axis of North America. Full-size DOI: 10.7717/peerj.9192/fig-19" figureDoi="http://doi.org/10.5281/zenodo.5825010" httpUri="https://zenodo.org/record/5825010/files/figure.png" pageId="55" pageNumber="56">Fig. 19</figureCitation>
|
||
), and stratigraphic position was ranked according to position within the lower, middle, and upper units of the Hell Creek Formation (HCF;
|
||
<tableCitation box="[1069,1170,1820,1847]" captionStart="Table 18" captionStartId="56.[525,580,986,1007]" captionTargetBox="[516,1525,1115,1471]" captionText="Table 18 Summary of raw and ranked data used in the Spearman correlation test of stratigraphic level in the Hell Creek Formation (and equivalents) and maturity for Tyrannosaurus rex. The specimens are organized by stratigraphic level and the values used in the correlation test are given in boldfaced columns; tied HCF ranks were converted to midranks. HCF, Hell Creek Formation." httpUri="http://table.plazi.org/id/DF5C84E2FFFCFF90FDEBFC523280FBFB" pageId="55" pageNumber="56" tableUuid="DF5C84E2FFFCFF90FDEBFC523280FBFB">Table 18</tableCitation>
|
||
;
|
||
<figureCitation box="[1185,1265,1820,1847]" captionStart="Figure 20" captionStartId="57.[524,589,675,697]" captionTargetBox="[502,1533,236,644]" captionTargetId="figure-369@57.[497,1123,233,649]" captionTargetPageId="57" captionText="Figure 20 Bivariate scatterplot showing the relationship between stratigraphic position with maturity among nine specimens of Tyrannosaurus rex. Growth stages (x-axis) and stratigraphic position (y-axis) have been converted to ranks. See Table 18 for the raw and ranked data. Maturity rank increases to the right; stratigraphic rank decreases from the origin (i.e., the upper HCF is closest to the origin, whereas the lower HCF is furthest from the origin). Full-size DOI: 10.7717/peerj.9192/fig-20" figureDoi="http://doi.org/10.5281/zenodo.5825012" httpUri="https://zenodo.org/record/5825012/files/figure.png" pageId="55" pageNumber="56">Fig. 20</figureCitation>
|
||
). To protect specimen localities and prevent trespassing, the precise locality coordinates are not provided here; locality data are available, to qualified researchers, from the museums where specimens are accessioned. Data for stratigraphic position was obtained from the literature (
|
||
<bibRefCitation author="Fowler DW" box="[623,771,1594,1620]" journalOrPublisher="PLOS ONE" pageId="56" pageNumber="57" pagination="e 0188426" part="12" refId="ref53571" refString="Fowler DW. 2017. Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America. PLOS ONE 12 (11): e 0188426 DOI 10.1371 / journal. pone. 0188426." title="Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America" type="journal article" year="2017">
|
||
<emphasis box="[623,771,1594,1620]" italics="true" pageId="56" pageNumber="57">Fowler, 2017</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gates TA & Gorscak E & Makovicky PJ" box="[786,1194,1594,1620]" journalOrPublisher="Journal of Paleontology" pageId="56" pageNumber="57" pagination="512 - 530" part="93" refId="ref53648" refString="Gates TA, Gorscak E, Makovicky PJ. 2019. New sharks and other chondrichthyans from the latest Maastrichtian (Late Cretaceous) of North America. Journal of Paleontology 93 (3): 512 - 530 DOI 10.1017 / jpa. 2018.92." title="New sharks and other chondrichthyans from the latest Maastrichtian (Late Cretaceous) of North America" type="journal article" year="2019">
|
||
<emphasis box="[786,1194,1594,1620]" italics="true" pageId="56" pageNumber="57">Gates, Gorscak & Makovicky, 2019</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Harrison WF & Nichols DJ & Henderson MD & Scherer RP" box="[1209,1449,1594,1620]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="56" pageNumber="57" pagination="239 - 250" refId="ref54085" refString="Harrison WF, Nichols DJ, Henderson MD, Scherer RP. 2013. Using pollen, leaves, and paleomagnetism to date a juvenile tyrannosaurid in upper Cretaceous rock. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 239 - 250." title="Using pollen, leaves, and paleomagnetism to date a juvenile tyrannosaurid in upper Cretaceous rock" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1209,1449,1594,1620]" italics="true" pageId="56" pageNumber="57">Harrison et al., 2013</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Horner JR & Goodwin MB & Myhrvold N." box="[498,924,1634,1660]" journalOrPublisher="PLOS ONE" pageId="56" pageNumber="57" pagination="e 16574" part="6" refId="ref54561" refString="Horner JR, Goodwin MB, Myhrvold N. 2011. Dinosaur census reveals abundant Tyrannosaurus rex and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA. PLOS ONE 6 (2): e 16574 DOI 10.1371 / journal. pone. 0016574." title="Dinosaur census reveals abundant Tyrannosaurus rex and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA" type="journal article" year="2011">
|
||
<emphasis box="[498,924,1634,1660]" italics="true" pageId="56" pageNumber="57">Horner, Goodwin & Myhrvold, 2011</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Leslie CE & Peppe DJ & Williamson TE & Heizler M & Jackson M & Atchley SC & Nordt L & Standhardt B." box="[939,1139,1634,1660]" journalOrPublisher="Geological Society of America Bulletin" pageId="56" pageNumber="57" pagination="1143 - 1163" part="130" refId="ref54871" refString="Leslie CE, Peppe DJ, Williamson TE, Heizler M, Jackson M, Atchley SC, Nordt L, Standhardt B. 2018. Revised age constraints for Late Cretaceous to early Paleocene terrestrial strata from the Dawson Creek section, Big Bend National Park, west Texas. Geological Society of America Bulletin 130 (7 - 8): 1143 - 1163 DOI 10.1130 / B 31785.1." title="Revised age constraints for Late Cretaceous to early Paleocene terrestrial strata from the Dawson Creek section, Big Bend National Park, west Texas" type="journal article" year="2018">
|
||
<emphasis box="[939,1139,1634,1660]" italics="true" pageId="56" pageNumber="57">Leslie et al., 2018</emphasis>
|
||
</bibRefCitation>
|
||
), aside from the specimen (
|
||
<materialsCitation collectionCode="DDM" pageId="56" pageNumber="57" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
) whose position was obtained from museum records.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FFF3FF9FFDEBFF70320DFE3F" httpUri="http://table.plazi.org/id/DF5C84E2FFF3FF9FFDEBFF70320DFE3F" pageId="55" pageNumber="56" startId="55.[525,580,248,269]" targetBox="[516,1525,407,1523]" targetIsTable="true" targetPageId="55">
|
||
<paragraph blockId="55.[525,1515,248,387]" pageId="55" pageNumber="56">
|
||
<emphasis bold="true" pageId="55" pageNumber="56">
|
||
Table 17 Summary of data used in the correlation test between geographic latitude and maturity of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[525,711,278,299]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="55" pageNumber="56" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[525,711,278,299]" italics="true" pageId="55" pageNumber="56">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
specimens.
|
||
</emphasis>
|
||
For each fossil, the columns are organized by specimen number, growth stage rank, geographic location rank, and level of resolution of the locality data. Specimens are listed by descending location rank (i.e., from north to south). Co, county; km, kilometer; m, meter; mi, miles; S, section; asterisk (
|
||
<emphasis box="[691,700,364,381]" italics="true" pageId="55" pageNumber="56">*</emphasis>
|
||
) indicates the resolution is almost certainly an underestimate.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="55" pageNumber="56">
|
||
<table box="[516,1525,407,1523]" gridcols="4" gridrows="30" pageId="55" pageNumber="56">
|
||
<tr box="[516,1525,407,429]" gridrow="0" pageId="55" pageNumber="56">
|
||
<th box="[516,709,407,429]" gridcol="0" gridrow="0" pageId="55" pageNumber="56">Specimen number</th>
|
||
<th box="[824,1006,407,429]" gridcol="1" gridrow="0" pageId="55" pageNumber="56">Maturity midrank</th>
|
||
<th box="[1121,1301,407,429]" gridcol="2" gridrow="0" pageId="55" pageNumber="56">Location midrank</th>
|
||
<th box="[1417,1525,407,429]" gridcol="3" gridrow="0" pageId="55" pageNumber="56">
|
||
<emphasis bold="true" box="[1417,1525,407,429]" pageId="55" pageNumber="56">Resolution</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1525,449,471]" gridrow="1" pageId="55" pageNumber="56">
|
||
<th box="[516,709,449,471]" gridcol="0" gridrow="1" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,709,449,470]" collectionCode="TMP" pageId="55" pageNumber="56" specimenCode="TMP 1981.006.0001">TMP 1981.012.0001</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,449,471]" gridcol="1" gridrow="1" pageId="55" pageNumber="56">12.5</td>
|
||
<td box="[1121,1301,449,471]" gridcol="2" gridrow="1" pageId="55" pageNumber="56">29</td>
|
||
<td box="[1417,1525,449,471]" gridcol="3" gridrow="1" pageId="55" pageNumber="56">km</td>
|
||
</tr>
|
||
<tr box="[516,1525,486,508]" gridrow="2" pageId="55" pageNumber="56">
|
||
<th box="[516,709,486,508]" gridcol="0" gridrow="2" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,709,486,508]" collectionCode="TMP" pageId="55" pageNumber="56" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,486,508]" gridcol="1" gridrow="2" pageId="55" pageNumber="56">9</td>
|
||
<td box="[1121,1301,486,508]" gridcol="2" gridrow="2" pageId="55" pageNumber="56">28</td>
|
||
<td box="[1417,1525,486,508]" gridcol="3" gridrow="2" pageId="55" pageNumber="56">km</td>
|
||
</tr>
|
||
<tr box="[516,1525,524,546]" gridrow="3" pageId="55" pageNumber="56">
|
||
<th box="[516,709,524,546]" gridcol="0" gridrow="3" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,632,524,545]" collectionCode="RSM" pageId="55" pageNumber="56" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,524,546]" gridcol="1" gridrow="3" pageId="55" pageNumber="56">5.5</td>
|
||
<td box="[1121,1301,524,546]" gridcol="2" gridrow="3" pageId="55" pageNumber="56">27</td>
|
||
<td box="[1417,1525,524,546]" gridcol="3" gridrow="3" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,561,583]" gridrow="4" pageId="55" pageNumber="56">
|
||
<th box="[516,709,561,583]" gridcol="0" gridrow="4" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,632,561,583]" collectionCode="RSM" pageId="55" pageNumber="56" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,561,583]" gridcol="1" gridrow="4" pageId="55" pageNumber="56">15</td>
|
||
<td box="[1121,1301,561,583]" gridcol="2" gridrow="4" pageId="55" pageNumber="56">26</td>
|
||
<td box="[1417,1525,561,583]" gridcol="3" gridrow="4" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,599,621]" gridrow="5" pageId="55" pageNumber="56">
|
||
<th box="[516,709,599,621]" gridcol="0" gridrow="5" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,606,599,620]" collectionCode="CM" pageId="55" pageNumber="56" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,599,621]" gridcol="1" gridrow="5" pageId="55" pageNumber="56">22.5</td>
|
||
<td box="[1121,1301,599,621]" gridcol="2" gridrow="5" pageId="55" pageNumber="56">25</td>
|
||
<td box="[1417,1525,599,621]" gridcol="3" gridrow="5" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,637,658]" gridrow="6" pageId="55" pageNumber="56">
|
||
<th box="[516,709,637,658]" gridcol="0" gridrow="6" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,646,637,658]" collectionCode="LACM" pageId="55" pageNumber="56" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,637,658]" gridcol="1" gridrow="6" pageId="55" pageNumber="56">1</td>
|
||
<td box="[1121,1301,637,658]" gridcol="2" gridrow="6" pageId="55" pageNumber="56">24</td>
|
||
<td box="[1417,1525,637,658]" gridcol="3" gridrow="6" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,674,696]" gridrow="7" pageId="55" pageNumber="56">
|
||
<th box="[516,709,674,696]" gridcol="0" gridrow="7" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,646,674,695]" collectionCode="LACM" pageId="55" pageNumber="56" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,674,696]" gridcol="1" gridrow="7" pageId="55" pageNumber="56">22.5</td>
|
||
<td box="[1121,1301,674,696]" gridcol="2" gridrow="7" pageId="55" pageNumber="56">22.5</td>
|
||
<td box="[1417,1525,674,696]" gridcol="3" gridrow="7" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,712,734]" gridrow="8" pageId="55" pageNumber="56">
|
||
<th box="[516,709,712,734]" gridcol="0" gridrow="8" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,646,712,734]" collectionCode="LACM" pageId="55" pageNumber="56" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,712,734]" gridcol="1" gridrow="8" pageId="55" pageNumber="56">7</td>
|
||
<td box="[1121,1301,712,734]" gridcol="2" gridrow="8" pageId="55" pageNumber="56">22.5</td>
|
||
<td box="[1417,1525,712,734]" gridcol="3" gridrow="8" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,750,771]" gridrow="9" pageId="55" pageNumber="56">
|
||
<th box="[516,709,750,771]" gridcol="0" gridrow="9" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,610,750,771]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,750,771]" gridcol="1" gridrow="9" pageId="55" pageNumber="56">20</td>
|
||
<td box="[1121,1301,750,771]" gridcol="2" gridrow="9" pageId="55" pageNumber="56">21</td>
|
||
<td box="[1417,1525,750,771]" gridcol="3" gridrow="9" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,787,809]" gridrow="10" pageId="55" pageNumber="56">
|
||
<th box="[516,709,787,809]" gridcol="0" gridrow="10" pageId="55" pageNumber="56">
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<materialsCitation box="[516,656,787,809]" collectionCode="UWBM" pageId="55" pageNumber="56" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
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||
</th>
|
||
<td box="[824,1006,787,809]" gridcol="1" gridrow="10" pageId="55" pageNumber="56">12.5</td>
|
||
<td box="[1121,1301,787,809]" gridcol="2" gridrow="10" pageId="55" pageNumber="56">20</td>
|
||
<td box="[1417,1525,787,809]" gridcol="3" gridrow="10" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,825,846]" gridrow="11" pageId="55" pageNumber="56">
|
||
<th box="[516,709,825,846]" gridcol="0" gridrow="11" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,622,825,846]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,825,846]" gridcol="1" gridrow="11" pageId="55" pageNumber="56">20</td>
|
||
<td box="[1121,1301,825,846]" gridcol="2" gridrow="11" pageId="55" pageNumber="56">19</td>
|
||
<td box="[1417,1525,825,846]" gridcol="3" gridrow="11" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,862,884]" gridrow="12" pageId="55" pageNumber="56">
|
||
<th box="[516,709,862,884]" gridcol="0" gridrow="12" pageId="55" pageNumber="56">
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||
<materialsCitation box="[516,622,862,884]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,862,884]" gridcol="1" gridrow="12" pageId="55" pageNumber="56">8</td>
|
||
<td box="[1121,1301,862,884]" gridcol="2" gridrow="12" pageId="55" pageNumber="56">18</td>
|
||
<td box="[1417,1525,862,884]" gridcol="3" gridrow="12" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,900,922]" gridrow="13" pageId="55" pageNumber="56">
|
||
<th box="[516,709,900,922]" gridcol="0" gridrow="13" pageId="55" pageNumber="56">
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||
<materialsCitation box="[516,610,900,922]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,900,922]" gridcol="1" gridrow="13" pageId="55" pageNumber="56">20</td>
|
||
<td box="[1121,1301,900,922]" gridcol="2" gridrow="13" pageId="55" pageNumber="56">17</td>
|
||
<td box="[1417,1525,900,922]" gridcol="3" gridrow="13" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,937,959]" gridrow="14" pageId="55" pageNumber="56">
|
||
<th box="[516,709,937,959]" gridcol="0" gridrow="14" pageId="55" pageNumber="56">
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<materialsCitation box="[516,660,938,959]" collectionCode="UCMP" pageId="55" pageNumber="56" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,937,959]" gridcol="1" gridrow="14" pageId="55" pageNumber="56">25</td>
|
||
<td box="[1121,1301,937,959]" gridcol="2" gridrow="14" pageId="55" pageNumber="56">16</td>
|
||
<td box="[1417,1525,937,959]" gridcol="3" gridrow="14" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,974,997]" gridrow="15" pageId="55" pageNumber="56">
|
||
<th box="[516,709,974,997]" gridcol="0" gridrow="15" pageId="55" pageNumber="56">
|
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<materialsCitation box="[516,610,975,996]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,974,997]" gridcol="1" gridrow="15" pageId="55" pageNumber="56">25</td>
|
||
<td box="[1121,1301,974,997]" gridcol="2" gridrow="15" pageId="55" pageNumber="56">15</td>
|
||
<td box="[1417,1525,974,997]" gridcol="3" gridrow="15" pageId="55" pageNumber="56">
|
||
m
|
||
<emphasis box="[1436,1445,974,991]" italics="true" pageId="55" pageNumber="56">*</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1013,1034]" gridrow="16" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1013,1034]" gridcol="0" gridrow="16" pageId="55" pageNumber="56">
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||
<materialsCitation box="[516,630,1013,1034]" collectionCode="DDM" pageId="55" pageNumber="56" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1013,1034]" gridcol="1" gridrow="16" pageId="55" pageNumber="56">2</td>
|
||
<td box="[1121,1301,1013,1034]" gridcol="2" gridrow="16" pageId="55" pageNumber="56">14</td>
|
||
<td box="[1417,1525,1013,1034]" gridcol="3" gridrow="16" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,1050,1072]" gridrow="17" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1050,1072]" gridcol="0" gridrow="17" pageId="55" pageNumber="56">
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<materialsCitation box="[516,658,1050,1072]" collectionCode="LACM" pageId="55" pageNumber="56" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1050,1072]" gridcol="1" gridrow="17" pageId="55" pageNumber="56">10</td>
|
||
<td box="[1121,1301,1050,1072]" gridcol="2" gridrow="17" pageId="55" pageNumber="56">13</td>
|
||
<td box="[1417,1525,1050,1072]" gridcol="3" gridrow="17" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,1088,1110]" gridrow="18" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1088,1110]" gridcol="0" gridrow="18" pageId="55" pageNumber="56">
|
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<materialsCitation box="[516,665,1088,1109]" collectionCode="BMRP" pageId="55" pageNumber="56" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1088,1110]" gridcol="1" gridrow="18" pageId="55" pageNumber="56">4</td>
|
||
<td box="[1121,1301,1088,1110]" gridcol="2" gridrow="18" pageId="55" pageNumber="56">12</td>
|
||
<td box="[1417,1525,1088,1110]" gridcol="3" gridrow="18" pageId="55" pageNumber="56">mi</td>
|
||
</tr>
|
||
<tr box="[516,1525,1125,1147]" gridrow="19" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1125,1147]" gridcol="0" gridrow="19" pageId="55" pageNumber="56">BMRP 2006.6.4</th>
|
||
<td box="[824,1006,1125,1147]" gridcol="1" gridrow="19" pageId="55" pageNumber="56">5.5</td>
|
||
<td box="[1121,1301,1125,1147]" gridcol="2" gridrow="19" pageId="55" pageNumber="56">11</td>
|
||
<td box="[1417,1525,1125,1147]" gridcol="3" gridrow="19" pageId="55" pageNumber="56">mi</td>
|
||
</tr>
|
||
<tr box="[516,1525,1163,1185]" gridrow="20" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1163,1185]" gridcol="0" gridrow="20" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,622,1163,1184]" collectionCode="MOR" pageId="55" pageNumber="56" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1163,1185]" gridcol="1" gridrow="20" pageId="55" pageNumber="56">27.5</td>
|
||
<td box="[1121,1301,1163,1185]" gridcol="2" gridrow="20" pageId="55" pageNumber="56">10</td>
|
||
<td box="[1417,1525,1163,1185]" gridcol="3" gridrow="20" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
<tr box="[516,1525,1201,1223]" gridrow="21" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1201,1223]" gridcol="0" gridrow="21" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,642,1201,1223]" collectionCode="CMNH" pageId="55" pageNumber="56" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1201,1223]" gridcol="1" gridrow="21" pageId="55" pageNumber="56">3</td>
|
||
<td box="[1121,1301,1201,1223]" gridcol="2" gridrow="21" pageId="55" pageNumber="56">9</td>
|
||
<td box="[1417,1525,1201,1223]" gridcol="3" gridrow="21" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,1238,1260]" gridrow="22" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1238,1260]" gridcol="0" gridrow="22" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,636,1238,1259]" collectionCode="UWGM" pageId="55" pageNumber="56" specimenCode="UWGM 181">UWGM 181</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1238,1260]" gridcol="1" gridrow="22" pageId="55" pageNumber="56">27.5</td>
|
||
<td box="[1121,1301,1238,1260]" gridcol="2" gridrow="22" pageId="55" pageNumber="56">8</td>
|
||
<td box="[1417,1525,1238,1260]" gridcol="3" gridrow="22" pageId="55" pageNumber="56">S</td>
|
||
</tr>
|
||
<tr box="[516,1525,1276,1298]" gridrow="23" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1276,1298]" gridcol="0" gridrow="23" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,667,1276,1297]" collectionCode="FMNH" pageId="55" pageNumber="56" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1276,1298]" gridcol="1" gridrow="23" pageId="55" pageNumber="56">29</td>
|
||
<td box="[1121,1301,1276,1298]" gridcol="2" gridrow="23" pageId="55" pageNumber="56">7</td>
|
||
<td box="[1417,1525,1276,1298]" gridcol="3" gridrow="23" pageId="55" pageNumber="56">mi</td>
|
||
</tr>
|
||
<tr box="[516,1525,1313,1335]" gridrow="24" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1313,1335]" gridcol="0" gridrow="24" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,641,1314,1335]" collectionCode="SDSM" pageId="55" pageNumber="56" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1313,1335]" gridcol="1" gridrow="24" pageId="55" pageNumber="56">15</td>
|
||
<td box="[1121,1301,1313,1335]" gridcol="2" gridrow="24" pageId="55" pageNumber="56">6</td>
|
||
<td box="[1417,1525,1313,1335]" gridcol="3" gridrow="24" pageId="55" pageNumber="56">mi</td>
|
||
</tr>
|
||
<tr box="[516,1525,1351,1373]" gridrow="25" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1351,1373]" gridcol="0" gridrow="25" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,707,1351,1373]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="55" pageNumber="56" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
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</th>
|
||
<td box="[824,1006,1351,1373]" gridcol="1" gridrow="25" pageId="55" pageNumber="56">17</td>
|
||
<td box="[1121,1301,1351,1373]" gridcol="2" gridrow="25" pageId="55" pageNumber="56">5</td>
|
||
<td box="[1417,1525,1351,1373]" gridcol="3" gridrow="25" pageId="55" pageNumber="56">Co</td>
|
||
</tr>
|
||
<tr box="[516,1525,1388,1410]" gridrow="26" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1388,1410]" gridcol="0" gridrow="26" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,606,1389,1410]" collectionCode="CM" pageId="55" pageNumber="56" specimenCode="CM 1400">CM 1400</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1388,1410]" gridcol="1" gridrow="26" pageId="55" pageNumber="56">11</td>
|
||
<td box="[1121,1301,1388,1410]" gridcol="2" gridrow="26" pageId="55" pageNumber="56">3.5</td>
|
||
<td box="[1417,1525,1388,1410]" gridcol="3" gridrow="26" pageId="55" pageNumber="56">Co</td>
|
||
</tr>
|
||
<tr box="[516,1525,1426,1448]" gridrow="27" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1426,1448]" gridcol="0" gridrow="27" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,674,1426,1448]" collectionCode="NHMUK" pageId="55" pageNumber="56" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1426,1448]" gridcol="1" gridrow="27" pageId="55" pageNumber="56">15</td>
|
||
<td box="[1121,1301,1426,1448]" gridcol="2" gridrow="27" pageId="55" pageNumber="56">3.5</td>
|
||
<td box="[1417,1525,1426,1448]" gridcol="3" gridrow="27" pageId="55" pageNumber="56">Co</td>
|
||
</tr>
|
||
<tr box="[516,1525,1464,1486]" gridrow="28" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1464,1486]" gridcol="0" gridrow="28" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,654,1464,1485]" collectionCode="UMNH" pageId="55" pageNumber="56" specimenCode="UMNH 11000">UMNH 11000</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1464,1486]" gridcol="1" gridrow="28" pageId="55" pageNumber="56">25</td>
|
||
<td box="[1121,1301,1464,1486]" gridcol="2" gridrow="28" pageId="55" pageNumber="56">2</td>
|
||
<td box="[1417,1525,1464,1486]" gridcol="3" gridrow="28" pageId="55" pageNumber="56">mi</td>
|
||
</tr>
|
||
<tr box="[516,1525,1501,1523]" gridrow="29" pageId="55" pageNumber="56">
|
||
<th box="[516,709,1501,1523]" gridcol="0" gridrow="29" pageId="55" pageNumber="56">
|
||
<materialsCitation box="[516,686,1501,1523]" collectionCode="NMMNH" pageId="55" pageNumber="56" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
</th>
|
||
<td box="[824,1006,1501,1523]" gridcol="1" gridrow="29" pageId="55" pageNumber="56">18</td>
|
||
<td box="[1121,1301,1501,1523]" gridcol="2" gridrow="29" pageId="55" pageNumber="56">1</td>
|
||
<td box="[1417,1525,1501,1523]" gridcol="3" gridrow="29" pageId="55" pageNumber="56">m</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825010" ID-Zenodo-Dep="5825010" httpUri="https://zenodo.org/record/5825010/files/figure.png" pageId="56" pageNumber="57" startId="56.[524,589,731,753]" targetBox="[497,1521,212,706]" targetPageId="56">
|
||
<paragraph blockId="56.[524,1515,731,900]" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" pageId="56" pageNumber="57">
|
||
Figure 19 Bivariate scatterplot showing the relationship between geographic location with maturity among 28 specimens
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[737,923,761,782]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="57" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[737,923,761,782]" italics="true" pageId="56" pageNumber="57">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages (
|
||
<emphasis box="[1086,1097,762,783]" italics="true" pageId="56" pageNumber="57">x</emphasis>
|
||
-axis) and geographic location (
|
||
<emphasis box="[1401,1411,762,783]" italics="true" pageId="56" pageNumber="57">y</emphasis>
|
||
-axis) have been converted to ranks. See Table 17 for the ranked data.
|
||
<emphasis box="[1115,1124,790,812]" italics="true" pageId="56" pageNumber="57">“</emphasis>
|
||
Montana North
|
||
<emphasis box="[1279,1288,790,812]" italics="true" pageId="56" pageNumber="57">”</emphasis>
|
||
refers to the region of Dawson, Garfield, and McCone counties, and
|
||
<emphasis box="[967,976,819,841]" italics="true" pageId="56" pageNumber="57">“</emphasis>
|
||
Montana South
|
||
<emphasis box="[1126,1135,819,841]" italics="true" pageId="56" pageNumber="57">”</emphasis>
|
||
refers to the region of Yellowstone and Carter counties. Maturity increases to the right along the
|
||
<emphasis box="[1063,1074,849,870]" italics="true" pageId="56" pageNumber="57">x</emphasis>
|
||
-axis; the
|
||
<emphasis box="[1165,1175,849,870]" italics="true" pageId="56" pageNumber="57">y</emphasis>
|
||
-axis follows the north-south axis of North America. Full-size DOI: 10.7717/peerj.9192/fig-19
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FFFCFF90FDEBFC523280FBFB" httpUri="http://table.plazi.org/id/DF5C84E2FFFCFF90FDEBFC523280FBFB" pageId="56" pageNumber="57" startId="56.[525,580,986,1007]" targetBox="[516,1525,1115,1471]" targetIsTable="true" targetPageId="56">
|
||
<paragraph blockId="56.[525,1516,985,1095]" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" pageId="56" pageNumber="57">
|
||
Table 18 Summary of raw and ranked data used in the Spearman correlation test of stratigraphic level in the Hell Creek Formation (and equivalents) and maturity for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1230,1417,1015,1036]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="56" pageNumber="57" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1230,1417,1015,1036]" italics="true" pageId="56" pageNumber="57">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The specimens are organized by stratigraphic level and the values used in the correlation test are given in boldfaced columns; tied HCF ranks were converted to midranks. HCF, Hell Creek Formation.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="56" pageNumber="57">
|
||
<table box="[516,1525,1115,1471]" gridcols="7" gridrows="9" pageId="56" pageNumber="57">
|
||
<tr box="[516,1525,1115,1167]" gridrow="0" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1115,1167]" gridcol="0" gridrow="0" pageId="56" pageNumber="57">Specimen</th>
|
||
<th box="[715,814,1115,1167]" gridcol="1" gridrow="0" pageId="56" pageNumber="57">HCF level</th>
|
||
<th box="[843,942,1115,1167]" gridcol="2" gridrow="0" pageId="56" pageNumber="57">HCF level rank</th>
|
||
<th box="[971,1072,1115,1167]" gridcol="3" gridrow="0" pageId="56" pageNumber="57">HCF rank</th>
|
||
<th box="[1101,1200,1115,1167]" gridcol="4" gridrow="0" pageId="56" pageNumber="57">HCF level midranks</th>
|
||
<th box="[1229,1363,1115,1167]" gridcol="5" gridrow="0" pageId="56" pageNumber="57">Growth stage</th>
|
||
<th box="[1391,1525,1115,1167]" gridcol="6" gridrow="0" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" pageId="56" pageNumber="57">Growth stage rank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1525,1186,1208]" gridrow="1" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1186,1208]" gridcol="0" gridrow="1" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,632,1186,1207]" collectionCode="RSM" pageId="56" pageNumber="57" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1186,1208]" gridcol="1" gridrow="1" pageId="56" pageNumber="57">Upper</td>
|
||
<td box="[843,942,1186,1208]" gridcol="2" gridrow="1" pageId="56" pageNumber="57">1</td>
|
||
<td box="[971,1072,1186,1208]" gridcol="3" gridrow="1" pageId="56" pageNumber="57">1</td>
|
||
<td box="[1101,1200,1186,1208]" gridcol="4" gridrow="1" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1186,1207]" pageId="56" pageNumber="57">2</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1186,1208]" gridcol="5" gridrow="1" pageId="56" pageNumber="57">5</td>
|
||
<td box="[1391,1525,1186,1208]" gridcol="6" gridrow="1" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1186,1207]" pageId="56" pageNumber="57">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1224,1246]" gridrow="2" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1224,1246]" gridcol="0" gridrow="2" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,632,1224,1246]" collectionCode="RSM" pageId="56" pageNumber="57" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1224,1246]" gridcol="1" gridrow="2" pageId="56" pageNumber="57">Upper</td>
|
||
<td box="[843,942,1224,1246]" gridcol="2" gridrow="2" pageId="56" pageNumber="57">1</td>
|
||
<td box="[971,1072,1224,1246]" gridcol="3" gridrow="2" pageId="56" pageNumber="57">2</td>
|
||
<td box="[1101,1200,1224,1246]" gridcol="4" gridrow="2" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1224,1245]" pageId="56" pageNumber="57">2</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1224,1246]" gridcol="5" gridrow="2" pageId="56" pageNumber="57">12</td>
|
||
<td box="[1391,1525,1224,1246]" gridcol="6" gridrow="2" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1224,1245]" pageId="56" pageNumber="57">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1261,1283]" gridrow="3" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1261,1283]" gridcol="0" gridrow="3" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,610,1261,1283]" collectionCode="MOR" pageId="56" pageNumber="57" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1261,1283]" gridcol="1" gridrow="3" pageId="56" pageNumber="57">Upper</td>
|
||
<td box="[843,942,1261,1283]" gridcol="2" gridrow="3" pageId="56" pageNumber="57">1</td>
|
||
<td box="[971,1072,1261,1283]" gridcol="3" gridrow="3" pageId="56" pageNumber="57">3</td>
|
||
<td box="[1101,1200,1261,1283]" gridcol="4" gridrow="3" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1261,1282]" pageId="56" pageNumber="57">2</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1261,1283]" gridcol="5" gridrow="3" pageId="56" pageNumber="57">17</td>
|
||
<td box="[1391,1525,1261,1283]" gridcol="6" gridrow="3" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1261,1282]" pageId="56" pageNumber="57">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1299,1321]" gridrow="4" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1299,1321]" gridcol="0" gridrow="4" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,630,1299,1320]" collectionCode="DDM" pageId="56" pageNumber="57" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1299,1321]" gridcol="1" gridrow="4" pageId="56" pageNumber="57">Middle</td>
|
||
<td box="[843,942,1299,1321]" gridcol="2" gridrow="4" pageId="56" pageNumber="57">2</td>
|
||
<td box="[971,1072,1299,1321]" gridcol="3" gridrow="4" pageId="56" pageNumber="57">4</td>
|
||
<td box="[1101,1200,1299,1321]" gridcol="4" gridrow="4" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1131,1299,1320]" pageId="56" pageNumber="57">4.5</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1299,1321]" gridcol="5" gridrow="4" pageId="56" pageNumber="57">2</td>
|
||
<td box="[1391,1525,1299,1321]" gridcol="6" gridrow="4" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1299,1320]" pageId="56" pageNumber="57">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1336,1359]" gridrow="5" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1336,1359]" gridcol="0" gridrow="5" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,665,1337,1358]" collectionCode="BMRP" pageId="56" pageNumber="57" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1336,1359]" gridcol="1" gridrow="5" pageId="56" pageNumber="57">Middle</td>
|
||
<td box="[843,942,1336,1359]" gridcol="2" gridrow="5" pageId="56" pageNumber="57">2</td>
|
||
<td box="[971,1072,1336,1359]" gridcol="3" gridrow="5" pageId="56" pageNumber="57">5</td>
|
||
<td box="[1101,1200,1336,1359]" gridcol="4" gridrow="5" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1131,1336,1357]" pageId="56" pageNumber="57">4.5</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1336,1359]" gridcol="5" gridrow="5" pageId="56" pageNumber="57">4</td>
|
||
<td box="[1391,1525,1336,1359]" gridcol="6" gridrow="5" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1336,1357]" pageId="56" pageNumber="57">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1374,1396]" gridrow="6" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1374,1396]" gridcol="0" gridrow="6" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,686,1374,1396]" collectionCode="NMMNH" pageId="56" pageNumber="57" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1374,1396]" gridcol="1" gridrow="6" pageId="56" pageNumber="57">Lower</td>
|
||
<td box="[843,942,1374,1396]" gridcol="2" gridrow="6" pageId="56" pageNumber="57">3</td>
|
||
<td box="[971,1072,1374,1396]" gridcol="3" gridrow="6" pageId="56" pageNumber="57">6</td>
|
||
<td box="[1101,1200,1374,1396]" gridcol="4" gridrow="6" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1374,1395]" pageId="56" pageNumber="57">7</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1374,1396]" gridcol="5" gridrow="6" pageId="56" pageNumber="57">15</td>
|
||
<td box="[1391,1525,1374,1396]" gridcol="6" gridrow="6" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1374,1395]" pageId="56" pageNumber="57">6</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1412,1434]" gridrow="7" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1412,1434]" gridcol="0" gridrow="7" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,622,1412,1434]" collectionCode="MOR" pageId="56" pageNumber="57" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1412,1434]" gridcol="1" gridrow="7" pageId="56" pageNumber="57">Lower</td>
|
||
<td box="[843,942,1412,1434]" gridcol="2" gridrow="7" pageId="56" pageNumber="57">3</td>
|
||
<td box="[971,1072,1412,1434]" gridcol="3" gridrow="7" pageId="56" pageNumber="57">7</td>
|
||
<td box="[1101,1200,1412,1434]" gridcol="4" gridrow="7" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1412,1433]" pageId="56" pageNumber="57">7</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1412,1434]" gridcol="5" gridrow="7" pageId="56" pageNumber="57">7</td>
|
||
<td box="[1391,1525,1412,1434]" gridcol="6" gridrow="7" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1412,1433]" pageId="56" pageNumber="57">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1449,1471]" gridrow="8" pageId="56" pageNumber="57">
|
||
<th box="[516,686,1449,1471]" gridcol="0" gridrow="8" pageId="56" pageNumber="57">
|
||
<materialsCitation box="[516,667,1449,1471]" collectionCode="FMNH" pageId="56" pageNumber="57" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
</th>
|
||
<td box="[715,814,1449,1471]" gridcol="1" gridrow="8" pageId="56" pageNumber="57">Lower</td>
|
||
<td box="[843,942,1449,1471]" gridcol="2" gridrow="8" pageId="56" pageNumber="57">3</td>
|
||
<td box="[971,1072,1449,1471]" gridcol="3" gridrow="8" pageId="56" pageNumber="57">8</td>
|
||
<td box="[1101,1200,1449,1471]" gridcol="4" gridrow="8" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1101,1113,1449,1470]" pageId="56" pageNumber="57">7</emphasis>
|
||
</td>
|
||
<td box="[1229,1363,1449,1471]" gridcol="5" gridrow="8" pageId="56" pageNumber="57">20</td>
|
||
<td box="[1391,1525,1449,1471]" gridcol="6" gridrow="8" pageId="56" pageNumber="57">
|
||
<emphasis bold="true" box="[1391,1403,1449,1470]" pageId="56" pageNumber="57">8</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="56.[498,1533,1739,1931]" box="[498,1003,1739,1767]" pageId="56" pageNumber="57">
|
||
<heading bold="true" box="[498,1003,1739,1767]" fontSize="11" level="3" pageId="56" pageNumber="57" reason="6">
|
||
<emphasis bold="true" box="[498,1003,1739,1767]" italics="true" pageId="56" pageNumber="57">Correlation with geographic position</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="56.[498,1533,1739,1931]" lastBlockId="57.[498,1530,870,1135]" lastPageId="57" lastPageNumber="58" pageId="56" pageNumber="57">
|
||
A Shapiro
|
||
<emphasis box="[617,632,1783,1808]" italics="true" pageId="56" pageNumber="57">–</emphasis>
|
||
Wilk test found that the ranked location data and growth rank data are normally distributed (
|
||
<emphasis box="[758,772,1823,1848]" italics="true" pageId="56" pageNumber="57">p</emphasis>
|
||
= 0.277 and 0.228, respectively). The geographic location test of ranked data (
|
||
<tableCitation box="[656,758,1862,1889]" captionStart="Table 17" captionStartId="55.[525,580,248,269]" captionTargetBox="[516,1525,407,1523]" captionText="Table 17 Summary of data used in the correlation test between geographic latitude and maturity of Tyrannosaurus rex specimens. For each fossil, the columns are organized by specimen number, growth stage rank, geographic location rank, and level of resolution of the locality data. Specimens are listed by descending location rank (i.e., from north to south). Co, county; km, kilometer; m, meter; mi, miles; S, section; asterisk (*) indicates the resolution is almost certainly an underestimate." httpUri="http://table.plazi.org/id/DF5C84E2FFF3FF9FFDEBFF70320DFE3F" pageId="56" pageNumber="57" tableUuid="DF5C84E2FFF3FF9FFDEBFF70320DFE3F">Table 17</tableCitation>
|
||
) resulted in a statistically nonsignificant (
|
||
<emphasis box="[1254,1268,1863,1888]" italics="true" pageId="56" pageNumber="57">p</emphasis>
|
||
= 0.219) correlation (
|
||
<emphasis box="[508,519,1903,1928]" italics="true" pageId="56" pageNumber="57">r</emphasis>
|
||
<subScript attach="left" box="[519,529,1914,1931]" fontSize="7" pageId="56" pageNumber="57">S</subScript>
|
||
= −0.236), indicating that the location of the specimens along the growth series was not influenced by geographic location. This result is robust given that it was possible to code a geographic location for nearly every specimen in the growth series (
|
||
<figureCitation box="[1385,1464,909,935]" captionStart="Figure 19" captionStartId="56.[524,589,731,753]" captionTargetBox="[497,1521,212,706]" captionTargetId="figure-133@56.[498,1458,234,704]" captionTargetPageId="56" captionText="Figure 19 Bivariate scatterplot showing the relationship between geographic location with maturity among 28 specimens Tyrannosaurus rex. Growth stages (x-axis) and geographic location (y-axis) have been converted to ranks. See Table 17 for the ranked data. “Montana North” refers to the region of Dawson, Garfield, and McCone counties, and “Montana South” refers to the region of Yellowstone and Carter counties. Maturity increases to the right along the x-axis; the y-axis follows the north-south axis of North America. Full-size DOI: 10.7717/peerj.9192/fig-19" figureDoi="http://doi.org/10.5281/zenodo.5825010" httpUri="https://zenodo.org/record/5825010/files/figure.png" pageId="57" pageNumber="58">Fig. 19</figureCitation>
|
||
). The correlation between unranked data (UTM coordinate northings) and growth stage was tested using a Pearson correlation test; data for 17 specimens were available for this test and the data are skewed to the right. The test resulted in a nonsignificant (
|
||
<emphasis box="[508,522,1070,1095]" italics="true" pageId="57" pageNumber="58">p</emphasis>
|
||
= 0.588) correlation coefficient (
|
||
<emphasis box="[912,923,1070,1095]" italics="true" pageId="57" pageNumber="58">r</emphasis>
|
||
= −0.141), which is consistent with the test on the ranked data.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825012" ID-Zenodo-Dep="5825012" httpUri="https://zenodo.org/record/5825012/files/figure.png" pageId="57" pageNumber="58" startId="57.[524,589,675,697]" targetBox="[502,1533,236,644]" targetPageId="57">
|
||
<paragraph blockId="57.[524,1515,675,815]" pageId="57" pageNumber="58">
|
||
<emphasis bold="true" pageId="57" pageNumber="58">
|
||
Figure 20 Bivariate scatterplot showing the relationship between stratigraphic position with maturity among nine specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[903,1094,705,726]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="57" pageNumber="58" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[903,1094,705,726]" italics="true" pageId="57" pageNumber="58">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stages (
|
||
<emphasis box="[1272,1283,705,726]" italics="true" pageId="57" pageNumber="58">x</emphasis>
|
||
-axis) and stratigraphic position (
|
||
<emphasis box="[616,626,735,756]" italics="true" pageId="57" pageNumber="58">y</emphasis>
|
||
-axis) have been converted to ranks. See Table 18 for the raw and ranked data. Maturity rank increases to the right; stratigraphic rank decreases from the origin (i.e., the upper HCF is closest to the origin, whereas the lower HCF is furthest from the origin). Full-size DOI: 10.7717/peerj.9192/fig-20
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="57.[498,1542,1174,1922]" box="[498,973,1174,1202]" pageId="57" pageNumber="58">
|
||
<heading bold="true" box="[498,973,1174,1202]" fontSize="11" level="3" pageId="57" pageNumber="58" reason="6">
|
||
<emphasis bold="true" box="[498,973,1174,1202]" italics="true" pageId="57" pageNumber="58">Correlation with stratigraphic level</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="57.[498,1542,1174,1922]" pageId="57" pageNumber="58">
|
||
The stratigraphic schema that is presented here for late Maastrichtian formations from western North America is largely based on
|
||
<bibRefCitation author="Fowler DW" box="[994,1152,1257,1283]" journalOrPublisher="PLOS ONE" pageId="57" pageNumber="58" pagination="e 0188426" part="12" refId="ref53571" refString="Fowler DW. 2017. Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America. PLOS ONE 12 (11): e 0188426 DOI 10.1371 / journal. pone. 0188426." title="Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America" type="journal article" year="2017">
|
||
<emphasis box="[994,1152,1257,1283]" italics="true" pageId="57" pageNumber="58">Fowler (2017)</emphasis>
|
||
</bibRefCitation>
|
||
, where the threefold stratigraphic division of the HCF of Montana is used as the basis of comparison with other units. As such, the comparison spans the last million years of the late Maastrichtian, which is divided into three chronostratigraphic slices, in descending order: 398 Kya (equivalent to chron 29r), ~286 Kya, and ~286 Kya (
|
||
<bibRefCitation author="Fowler DW" box="[984,1132,1417,1443]" journalOrPublisher="PLOS ONE" pageId="57" pageNumber="58" pagination="e 0188426" part="12" refId="ref53571" refString="Fowler DW. 2017. Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America. PLOS ONE 12 (11): e 0188426 DOI 10.1371 / journal. pone. 0188426." title="Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America" type="journal article" year="2017">
|
||
<emphasis box="[984,1132,1417,1443]" italics="true" pageId="57" pageNumber="58">Fowler, 2017</emphasis>
|
||
</bibRefCitation>
|
||
). Using this approach, the Frenchman Formation of Saskatchewan is equivalent to the upper HCF (
|
||
<bibRefCitation author="Fowler DW" box="[1361,1509,1457,1483]" journalOrPublisher="PLOS ONE" pageId="57" pageNumber="58" pagination="e 0188426" part="12" refId="ref53571" refString="Fowler DW. 2017. Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America. PLOS ONE 12 (11): e 0188426 DOI 10.1371 / journal. pone. 0188426." title="Revised geochronology, correlation, and dinosaur stratigraphic ranges of the Santonian-Maastrichtian (Late Cretaceous) formations of the Western Interior of North America" type="journal article" year="2017">
|
||
<emphasis box="[1361,1509,1457,1483]" italics="true" pageId="57" pageNumber="58">Fowler, 2017</emphasis>
|
||
</bibRefCitation>
|
||
). The radiometric dates and stratigraphic revisions of
|
||
<bibRefCitation author="Leslie CE & Peppe DJ & Williamson TE & Heizler M & Jackson M & Atchley SC & Nordt L & Standhardt B." box="[1117,1329,1496,1523]" journalOrPublisher="Geological Society of America Bulletin" pageId="57" pageNumber="58" pagination="1143 - 1163" part="130" refId="ref54871" refString="Leslie CE, Peppe DJ, Williamson TE, Heizler M, Jackson M, Atchley SC, Nordt L, Standhardt B. 2018. Revised age constraints for Late Cretaceous to early Paleocene terrestrial strata from the Dawson Creek section, Big Bend National Park, west Texas. Geological Society of America Bulletin 130 (7 - 8): 1143 - 1163 DOI 10.1130 / B 31785.1." title="Revised age constraints for Late Cretaceous to early Paleocene terrestrial strata from the Dawson Creek section, Big Bend National Park, west Texas" type="journal article" year="2018">
|
||
<emphasis box="[1117,1329,1496,1523]" italics="true" pageId="57" pageNumber="58">Leslie et al. (2018)</emphasis>
|
||
</bibRefCitation>
|
||
were followed to correlate the Tornillo Formation of Texas with the upper HCF. Given the imprecise dating of the Scollard and Willow Creek Formations of Alberta, and ambiguous correlation between them and the upper and middle members of the HCF, specimens from those units were excluded. This comparison was made aware of the fact that the stratigraphic position of specimens might be affected by time transgression and so might not be equivalent. Resolving that issue is beyond the scope of this work, the results of which are offered here as a hypothesis for further, more rigorous testing of stratigraphic correlation.
|
||
</paragraph>
|
||
<paragraph blockId="57.[498,1542,1174,1922]" lastBlockId="58.[498,1542,234,579]" lastPageId="58" lastPageNumber="59" pageId="57" pageNumber="58">
|
||
A Shapiro
|
||
<emphasis box="[649,664,1816,1841]" italics="true" pageId="57" pageNumber="58">–</emphasis>
|
||
Wilk test found that the ranked stratigraphic data are not normally distributed (
|
||
<emphasis box="[642,656,1856,1881]" italics="true" pageId="57" pageNumber="58">p</emphasis>
|
||
= 0.030), whereas the growth rank data are normally distributed (
|
||
<emphasis box="[1419,1433,1856,1881]" italics="true" pageId="57" pageNumber="58">p</emphasis>
|
||
= 0.933). A Spearman rank correlation test was run, which recovered a nonsignificant (
|
||
<emphasis box="[1417,1431,1896,1921]" italics="true" pageId="57" pageNumber="58">p</emphasis>
|
||
= 0.654) correlation (
|
||
<emphasis box="[644,655,235,260]" italics="true" pageId="58" pageNumber="59">r</emphasis>
|
||
<subScript attach="left" box="[655,665,246,263]" fontSize="7" pageId="58" pageNumber="59">S</subScript>
|
||
= 0.189) between stratigraphic position and maturity (
|
||
<figureCitation box="[1316,1395,234,260]" captionStart="Figure 20" captionStartId="57.[524,589,675,697]" captionTargetBox="[502,1533,236,644]" captionTargetId="figure-369@57.[497,1123,233,649]" captionTargetPageId="57" captionText="Figure 20 Bivariate scatterplot showing the relationship between stratigraphic position with maturity among nine specimens of Tyrannosaurus rex. Growth stages (x-axis) and stratigraphic position (y-axis) have been converted to ranks. See Table 18 for the raw and ranked data. Maturity rank increases to the right; stratigraphic rank decreases from the origin (i.e., the upper HCF is closest to the origin, whereas the lower HCF is furthest from the origin). Full-size DOI: 10.7717/peerj.9192/fig-20" figureDoi="http://doi.org/10.5281/zenodo.5825012" httpUri="https://zenodo.org/record/5825012/files/figure.png" pageId="58" pageNumber="59">Fig. 20</figureCitation>
|
||
;
|
||
<tableCitation box="[1410,1512,234,260]" captionStart="Table 18" captionStartId="56.[525,580,986,1007]" captionTargetBox="[516,1525,1115,1471]" captionText="Table 18 Summary of raw and ranked data used in the Spearman correlation test of stratigraphic level in the Hell Creek Formation (and equivalents) and maturity for Tyrannosaurus rex. The specimens are organized by stratigraphic level and the values used in the correlation test are given in boldfaced columns; tied HCF ranks were converted to midranks. HCF, Hell Creek Formation." httpUri="http://table.plazi.org/id/DF5C84E2FFFCFF90FDEBFC523280FBFB" pageId="58" pageNumber="59" tableUuid="DF5C84E2FFFCFF90FDEBFC523280FBFB">Table 18</tableCitation>
|
||
). However, this result is not robust given that only eight specimens are included in the sample and only two are from the middle member of the HCF (BMRP 2002.4.1;
|
||
<bibRefCitation author="Harrison WF & Nichols DJ & Henderson MD & Scherer RP" box="[498,738,354,380]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="58" pageNumber="59" pagination="239 - 250" refId="ref54085" refString="Harrison WF, Nichols DJ, Henderson MD, Scherer RP. 2013. Using pollen, leaves, and paleomagnetism to date a juvenile tyrannosaurid in upper Cretaceous rock. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 239 - 250." title="Using pollen, leaves, and paleomagnetism to date a juvenile tyrannosaurid in upper Cretaceous rock" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[498,738,354,380]" italics="true" pageId="58" pageNumber="59">Harrison et al., 2013</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<materialsCitation box="[752,894,354,380]" collectionCode="DDM" pageId="58" pageNumber="59" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
) (
|
||
<figureCitation box="[920,999,354,380]" captionStart="Figure 20" captionStartId="57.[524,589,675,697]" captionTargetBox="[502,1533,236,644]" captionTargetId="figure-369@57.[497,1123,233,649]" captionTargetPageId="57" captionText="Figure 20 Bivariate scatterplot showing the relationship between stratigraphic position with maturity among nine specimens of Tyrannosaurus rex. Growth stages (x-axis) and stratigraphic position (y-axis) have been converted to ranks. See Table 18 for the raw and ranked data. Maturity rank increases to the right; stratigraphic rank decreases from the origin (i.e., the upper HCF is closest to the origin, whereas the lower HCF is furthest from the origin). Full-size DOI: 10.7717/peerj.9192/fig-20" figureDoi="http://doi.org/10.5281/zenodo.5825012" httpUri="https://zenodo.org/record/5825012/files/figure.png" pageId="58" pageNumber="59">Fig. 20</figureCitation>
|
||
). Therefore, more specimen data are required for a rigorous test of the noncorrelation between maturity and stratigraphic position. Regardless, the scatterplot shows that a young adult, two adults, and senescent adult have been collected from the lower member of the HCF, two juveniles from the middle unit, and one subadult and two adults from the upper member; that is, broadly overlapping growth series have been recovered throughout the unit.
|
||
</paragraph>
|
||
<paragraph blockId="58.[498,1542,624,1932]" box="[498,1221,624,654]" pageId="58" pageNumber="59">
|
||
<heading bold="true" box="[498,1221,624,654]" fontSize="12" level="2" pageId="58" pageNumber="59" reason="0">
|
||
<emphasis bold="true" box="[498,1221,624,654]" pageId="58" pageNumber="59">
|
||
Dimorphism hypothesis (sensu
|
||
<bibRefCitation author="Carpenter K." box="[979,1211,624,654]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis bold="true" box="[979,1211,624,654]" italics="true" pageId="58" pageNumber="59">Carpenter, 1990</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="58.[498,1542,624,1932]" pageId="58" pageNumber="59">
|
||
<bibRefCitation author="Carpenter K." box="[498,696,670,696]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[498,696,670,696]" italics="true" pageId="58" pageNumber="59">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
identified two sets of characters in the skull and postcranium of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1467,1536,671,696]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="58" pageNumber="59" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1467,1536,671,696]" italics="true" pageId="58" pageNumber="59">T. rex</emphasis>
|
||
</taxonomicName>
|
||
that he considered to represent patternless variation (noise; i.e., individual variation) on the one hand, and patterned variation (signal; i.e., sexual variation) on the other. Noise included: maxilla depth, size of the maxillary fenestra, shape of the maxillary fenestra, size of the antorbital fenestra, shape of the antorbital fenestra, position of the lacrimal process of the maxilla, position of the jugal process of the maxilla, the shape of the jugal process of the maxilla, and shape of the dentary. In contrast, signal included: robust and gracile morphs of cervical vertebrae and angle of the ischium from the caudal series. The cladistic results obtained here agree that, among adults, the depth of the maxilla and size and shape of the maxillary and antorbital fenestrae, the position and shape of the lacrimal and jugal processes of the maxilla, and shape of the dentary, are noise.
|
||
</paragraph>
|
||
<paragraph blockId="58.[498,1542,624,1932]" pageId="58" pageNumber="59">
|
||
Also,
|
||
<bibRefCitation author="Carpenter K." box="[597,793,1108,1135]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[597,793,1108,1135]" italics="true" pageId="58" pageNumber="59">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
found that the maxilla
|
||
<materialsCitation box="[1067,1239,1108,1135]" collectionCode="TMM" pageId="58" pageNumber="59" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
|
||
was an outlier in contrast to the sample of adult maxillae, which were scaled to the rostral end of the maxilla and the rostral end of the maxillary fenestra. The incomplete and small
|
||
<materialsCitation box="[1297,1470,1188,1215]" collectionCode="TMM" pageId="58" pageNumber="59" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
|
||
was scaled to the height of the maxillary fenestra (
|
||
<bibRefCitation author="Carpenter K." box="[1042,1228,1228,1254]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[1042,1228,1228,1254]" italics="true" pageId="58" pageNumber="59">Carpenter, 1990</emphasis>
|
||
</bibRefCitation>
|
||
), which is proportionately taller than in adults (e.g.,
|
||
<materialsCitation box="[802,1041,1267,1294]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="58" pageNumber="59" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
). The comparison showed that it is different from the others in that the rostral margin of the bone falls short of the adults, which was used by
|
||
<bibRefCitation author="Carpenter K." box="[728,926,1347,1374]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[728,926,1347,1374]" italics="true" pageId="58" pageNumber="59">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
to argue the specimen represents a new taxon; however, based on the results found here, this difference almost certainly arises from the fact its teeth are not as enlarged in adults and the internal sinuses are not expanded, changes that greatly reshape the bone in adults. Therefore, the difference in shape is most simply explained by maturity; when included in the cladistic analysis of ontogeny, the specimen falls out in a polytomy that includes subadult, young adult, and adult specimens (
|
||
<figureCitation box="[508,591,1586,1613]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="58" pageNumber="59">Fig. 1B</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="58.[498,1542,624,1932]" lastBlockId="59.[498,1542,234,1337]" lastPageId="59" lastPageNumber="60" pageId="58" pageNumber="59">
|
||
<bibRefCitation author="Carpenter K." box="[530,728,1627,1653]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[530,728,1627,1653]" italics="true" pageId="58" pageNumber="59">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
found, qualitatively, that the cervical series of
|
||
<materialsCitation box="[1280,1513,1626,1653]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="58" pageNumber="59" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
was less massive than that of
|
||
<materialsCitation box="[848,1044,1666,1693]" collectionCode="NHMUK" pageId="58" pageNumber="59" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
. In particular,
|
||
<bibRefCitation author="Carpenter K." box="[1218,1416,1666,1693]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="58" pageNumber="59" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[1218,1416,1666,1693]" italics="true" pageId="58" pageNumber="59">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
drew attention to the form of the atlantal intercentrum and that of the spinous process of the axis and the third cervical vertebra, characters that were included in the cladistic analysis presented here (characters 1,210, 1,256, and 1,280, respectively; see Data S1). The results found that those specimens are ontogenetically sequential, where
|
||
<materialsCitation box="[1272,1505,1825,1852]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="58" pageNumber="59" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
is less mature than
|
||
<materialsCitation box="[700,894,1866,1892]" collectionCode="NHMUK" pageId="58" pageNumber="59" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
, indicating that the differences (slender to massive) are ontogenetic. It is worth pointing out that the illustrations that were used to show the differences between the specimens are mislabeled, where the massive
|
||
<materialsCitation box="[1318,1511,234,261]" collectionCode="NHMUK" pageId="59" pageNumber="60" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
is mistakenly labeled as the slender
|
||
<materialsCitation box="[894,1129,274,301]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="59" pageNumber="60" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
, and vice versa (
|
||
<bibRefCitation author="Carpenter K." box="[1328,1513,274,301]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[1328,1513,274,301]" italics="true" pageId="59" pageNumber="60">Carpenter, 1990</emphasis>
|
||
</bibRefCitation>
|
||
:
|
||
<figureCitation box="[498,699,314,340]" captionStart="Figure 10" captionStartId="21.[524,589,1489,1511]" captionTargetBox="[503,1531,1030,1457]" captionTargetId="figure-217@21.[497,1128,1026,1462]" captionTargetPageId="21" captionText="Figure 10 The frequency distribution of postcranial synontomorphies in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. Changes to the appendicular skeleton dominate in the transition between juvenile and subadult, whereas changes to the pelvic girdle and axial skeleton occur late in adulthood. Full-size DOI: 10.7717/peerj.9192/fig-10" figureDoi="http://doi.org/10.5281/zenodo.5824992" httpUri="https://zenodo.org/record/5824992/files/figure.png" pageId="59" pageNumber="60">Fig. 10.4A and B</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="59.[498,1542,234,1337]" pageId="59" pageNumber="60">
|
||
Finally,
|
||
<bibRefCitation author="Carpenter K." box="[625,823,354,380]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[625,823,354,380]" italics="true" pageId="59" pageNumber="60">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
compared the several features of the ischium between three specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[659,728,394,419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="59" pageNumber="60" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[659,728,394,419]" italics="true" pageId="59" pageNumber="60">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. He identified the orientation of the ischia as informative, where in two (
|
||
<materialsCitation box="[558,672,434,460]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
, TMP 1981.006.0001) the bone extends sharply posteroventrally whereas the third (
|
||
<materialsCitation box="[720,958,473,500]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="59" pageNumber="60" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
) extends at a lower angle.
|
||
<bibRefCitation author="Carpenter K." box="[1274,1471,473,500]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[1274,1471,473,500]" italics="true" pageId="59" pageNumber="60">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
hypothesized that the steep condition was required for passing eggs in life, and the bearers were female, whereas the bearer of the other form was male. The results of the cladistic analysis here found that of these three specimens,
|
||
<materialsCitation box="[1086,1321,593,619]" collectionCode="TMP" pageId="59" pageNumber="60" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
is the least mature and that
|
||
<materialsCitation box="[606,715,633,659]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
is the most mature. Therefore, a growth sequence from steep (TMP 1981.006.0001) to less steep (
|
||
<materialsCitation box="[916,1154,672,699]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="59" pageNumber="60" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
), and back to steep (
|
||
<materialsCitation box="[1394,1508,673,699]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
) is seen. If the angles of divergence are real, then a lack of signal (noise) is probably the best explanation for the observation.
|
||
</paragraph>
|
||
<paragraph blockId="59.[498,1542,234,1337]" pageId="59" pageNumber="60">
|
||
However, the position of the tip of the ischium is an important landmark for this comparison, but it is missing from
|
||
<materialsCitation box="[911,1020,832,858]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Osborn HF" box="[1038,1190,832,858]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="59" pageNumber="60" pagination="281 - 296" part="22" refId="ref55443" refString="Osborn HF. 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication). Bulletin of the American Museum of Natural History 22: 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication)" type="journal article" year="1906">
|
||
<emphasis box="[1038,1190,832,858]" italics="true" pageId="59" pageNumber="60">Osborn, 1906</emphasis>
|
||
</bibRefCitation>
|
||
: fig. 7) whereas it is shown as complete in
|
||
<bibRefCitation author="Carpenter K." box="[644,830,872,898]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[644,830,872,898]" italics="true" pageId="59" pageNumber="60">Carpenter (1990</emphasis>
|
||
</bibRefCitation>
|
||
: fig. 10.5). The complete condition presumably reflects restoration of the bone as indicated by the presence of a small boot at its tip (
|
||
<bibRefCitation author="Carpenter K." editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis italics="true" pageId="59" pageNumber="60">Carpenter, 1990</emphasis>
|
||
</bibRefCitation>
|
||
: fig. 10.5), a structure that is not seen in complete specimens of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1309,1377,952,977]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="59" pageNumber="60" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1309,1377,952,977]" italics="true" pageId="59" pageNumber="60">T. rex</emphasis>
|
||
</taxonomicName>
|
||
or in tyrannosaurids in general. Also, the shaft of the ischium in
|
||
<materialsCitation box="[1194,1304,992,1018]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
is intermediate in position between the two other specimens in that it overlaps the dorsal margin of the ischium of
|
||
<materialsCitation box="[625,855,1071,1098]" collectionCode="TMP" pageId="59" pageNumber="60" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Carpenter K." box="[871,1052,1071,1098]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[871,1052,1071,1098]" italics="true" pageId="59" pageNumber="60">Carpenter, 1990</emphasis>
|
||
</bibRefCitation>
|
||
: fig. 10.5). Given these issues, the character was included a posteriori in the cladistic data matrix and only the three specimens mentioned by
|
||
<bibRefCitation author="Carpenter K." box="[668,864,1151,1177]" editor="Carpenter K & Currie PJ" journalOrPublisher="University of Cambridge Press" pageId="59" pageNumber="60" pagination="141 - 145" refId="ref52602" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. In: Carpenter K, Currie PJ, eds. Dinosaur Systematics: Approaches and Perspective. University of Cambridge Press, 141 - 145." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics: Approaches and Perspective" year="1990">
|
||
<emphasis box="[668,864,1151,1177]" italics="true" pageId="59" pageNumber="60">Carpenter (1990)</emphasis>
|
||
</bibRefCitation>
|
||
were coded in the analysis, using the state assignments given by him. The analysis recovered one 3,054-step ontogram and the topology is unaffected. Therefore, based on this sample, ischial divergence does not group specimens together, indicating that it is not dimorphic; regardless, a larger data set for this character is required for a rigorous test of whether the two states are valid to begin with.
|
||
</paragraph>
|
||
<paragraph blockId="59.[498,1542,1381,1932]" box="[498,1506,1381,1412]" pageId="59" pageNumber="60">
|
||
<heading bold="true" box="[498,1506,1381,1412]" fontSize="12" level="2" pageId="59" pageNumber="60" reason="0">
|
||
<emphasis bold="true" box="[498,1506,1381,1412]" pageId="59" pageNumber="60">
|
||
Sexual dimorphism and “
|
||
<emphasis bold="true" box="[881,1150,1381,1412]" italics="true" pageId="59" pageNumber="60">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[881,1106,1381,1411]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="59" pageNumber="60" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
“x
|
||
</emphasis>
|
||
”” (sensu
|
||
<bibRefCitation author="Larson P." box="[1302,1496,1381,1411]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="59" pageNumber="60" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis bold="true" box="[1302,1496,1381,1411]" italics="true" pageId="59" pageNumber="60">Larson, 2008</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="59.[498,1542,1381,1932]" pageId="59" pageNumber="60">
|
||
A topological pattern of sexual dimorphism was not recovered in the ontogram, which is linear and pectinate (
|
||
<figureCitation box="[750,816,1467,1493]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="59" pageNumber="60">Fig. 2</figureCitation>
|
||
). If dimorphism was present in the data, which covers the entire skeleton, and if the sample included examples of each sex, then a branch of females that includes
|
||
<materialsCitation box="[750,933,1546,1573]" collectionCode="BMRP" pageId="59" pageNumber="60" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
and
|
||
<materialsCitation box="[991,1122,1546,1573]" collectionCode="MOR" pageId="59" pageNumber="60" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, the only purportedly unambiguous females in the sample, should be united by the presence of shared skeletal sexual correlates, aside from femoral medullary bone (
|
||
<bibRefCitation author="Schweitzer MH & Wittmeyer JL & Horner JR" box="[1060,1513,1626,1653]" journalOrPublisher="Science" pageId="59" pageNumber="60" pagination="1456 - 1460" part="308" refId="ref55976" refString="Schweitzer MH, Wittmeyer JL, Horner JR. 2005. Gender-specific reproductive tissue in ratites and Tyrannosaurus rex. Science 308 (5727): 1456 - 1460 DOI 10.1126 / science. 1112158." title="Gender-specific reproductive tissue in ratites and Tyrannosaurus rex" type="journal article" year="2005">
|
||
<emphasis box="[1060,1513,1626,1653]" italics="true" pageId="59" pageNumber="60">Schweitzer, Wittmeyer & Horner, 2005</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[498,763,1666,1692]" journalOrPublisher="Science Advances" pageId="59" pageNumber="60" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[498,763,1666,1692]" italics="true" pageId="59" pageNumber="60">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
). The failure to recover a female branch indicates that such correlates are absent from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[806,873,1707,1732]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="59" pageNumber="60" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[806,873,1707,1732]" italics="true" pageId="59" pageNumber="60">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Neither did a comparison of individual variation recover a dimorphic pattern (see above). The alternative explanation for these results is that
|
||
<materialsCitation box="[498,680,1786,1812]" collectionCode="BMRP" pageId="59" pageNumber="60" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
and
|
||
<materialsCitation box="[739,868,1786,1813]" collectionCode="MOR" pageId="59" pageNumber="60" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
are the only females in the sample, but this is improbable given the high number of adult-grade specimens in the analysis.
|
||
</paragraph>
|
||
<paragraph blockId="59.[498,1542,1381,1932]" lastBlockId="60.[498,1542,1268,1893]" lastPageId="60" lastPageNumber="61" pageId="59" pageNumber="60">
|
||
Regardless, several specimens included in this study have been hypothesized as male (
|
||
<materialsCitation box="[506,744,1905,1931]" collectionCode="TMP" pageId="59" pageNumber="60" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
,
|
||
<materialsCitation box="[754,870,1905,1931]" collectionCode="MOR" pageId="59" pageNumber="60" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation box="[880,996,1905,1932]" collectionCode="MOR" pageId="59" pageNumber="60" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
) or female (
|
||
<materialsCitation box="[1133,1265,1905,1932]" collectionCode="MOR" pageId="59" pageNumber="60" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1275,1418,1905,1932]" collectionCode="RSM" pageId="59" pageNumber="60" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1428,1537,1905,1931]" collectionCode="CM" pageId="59" pageNumber="60" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,684,1269,1295]" collectionCode="FMNH" pageId="60" pageNumber="61" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
) and so an account of the difference between those results (
|
||
<bibRefCitation author="Larson P." box="[1383,1532,1269,1295]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[1383,1532,1269,1295]" italics="true" pageId="60" pageNumber="61">Larson, 2008</emphasis>
|
||
</bibRefCitation>
|
||
) and those obtained here is required. No grouping pattern is seen when male (
|
||
<emphasis box="[1383,1394,1308,1334]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
gracile
|
||
<emphasis box="[1470,1481,1308,1334]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
) and female (
|
||
<emphasis box="[594,605,1348,1374]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
robust
|
||
<emphasis box="[680,691,1348,1374]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
) morphs (sensu
|
||
<bibRefCitation author="Larson P." box="[892,1042,1348,1374]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[892,1042,1348,1374]" italics="true" pageId="60" pageNumber="61">Larson, 2008</emphasis>
|
||
</bibRefCitation>
|
||
) are mapped onto the ontogram, where both
|
||
<emphasis box="[561,573,1388,1414]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
morphs
|
||
<emphasis box="[662,674,1388,1414]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
are interspersed among the adult specimens (
|
||
<figureCitation box="[1219,1298,1388,1414]" captionStart="Figure 21" captionStartId="60.[524,589,957,979]" captionTargetBox="[498,1541,234,945]" captionTargetId="graphics-291@60.[498,1270,245,922]" captionTargetPageId="60" captionText="Figure 21 Sex dimorphs and the taxon “Tyrannosaurus “x”” of Larson (2008) mapped onto the ontogram of Tyrannosaurus rex. A transitional pattern is not seen between gracile and robust morphs; if sexual dimorphism was present, then the “gracile” and “robust” morphs should group along separate branches, which is not seen. Also, specimens referred to the taxon “T. “x”” do not form a clade, indicating that it is not a valid taxon. The pattern seen here is what is expected for a species without sexual dimorphism. Specimens considered in Larson (2008) as gracile are in boldface italics with a boldface “G”; specimens considered in Larson (2008) as robust are in boldface with a boldface “R”; specimens considered in Larson (2008) as referable to “T. “x”” are in italics and marked with an “X”. Full-size DOI: 10.7717/peerj.9192/fig-21" figureDoi="http://doi.org/10.5281/zenodo.5825014" httpUri="https://zenodo.org/record/5825014/files/figure.png" pageId="60" pageNumber="61">Fig. 21</figureCitation>
|
||
). A biologically meaningful signal would be indicated if, for example, gracile morphs occurred earlier in growth than the robust morphs, or vice versa. Given that both morphs are seen in young adult and adult growth stages, the simplest explanation is that they do not represent a developmental pattern aside from individual variation upon the attainment of the asymptote of size.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825014" ID-Zenodo-Dep="5825014" httpUri="https://zenodo.org/record/5825014/files/figure.png" pageId="60" pageNumber="61" startId="60.[524,589,957,979]" targetBox="[498,1541,234,945]" targetPageId="60">
|
||
<paragraph blockId="60.[524,1516,957,1213]" pageId="60" pageNumber="61">
|
||
<emphasis bold="true" box="[524,927,957,979]" pageId="60" pageNumber="61">Figure 21 Sex dimorphs and the taxon</emphasis>
|
||
“
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[948,1097,958,979]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="60" pageNumber="61" phylum="Chordata" rank="genus">
|
||
<emphasis bold="true" box="[948,1097,958,979]" italics="true" pageId="60" pageNumber="61">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
“
|
||
<emphasis bold="true" box="[1116,1127,958,979]" italics="true" pageId="60" pageNumber="61">x</emphasis>
|
||
””
|
||
<emphasis bold="true" pageId="60" pageNumber="61">
|
||
of
|
||
<bibRefCitation author="Larson P." box="[1188,1329,957,979]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis bold="true" box="[1188,1329,957,979]" italics="true" pageId="60" pageNumber="61">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
mapped onto the ontogram of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[668,860,987,1008]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="60" pageNumber="61" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[668,860,987,1008]" italics="true" pageId="60" pageNumber="61">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
A transitional pattern is not seen between gracile and robust morphs; if sexual dimorphism was present, then the
|
||
<emphasis box="[1030,1039,1016,1038]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
gracile
|
||
<emphasis box="[1102,1111,1016,1038]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
and
|
||
<emphasis box="[1161,1170,1016,1038]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
robust
|
||
<emphasis box="[1232,1241,1016,1038]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
morphs should group along separate branches, which is not seen. Also, specimens referred to the taxon
|
||
<emphasis box="[1242,1265,1045,1067]" italics="true" pageId="60" pageNumber="61">“T</emphasis>
|
||
.
|
||
<emphasis box="[1277,1317,1045,1067]" italics="true" pageId="60" pageNumber="61">“x””</emphasis>
|
||
do not form a clade, indicating that it is not a valid taxon. The pattern seen here is what is expected for a species without sexual dimorphism. Specimens considered in
|
||
<bibRefCitation author="Larson P." box="[893,1025,1104,1125]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[893,1025,1104,1125]" italics="true" pageId="60" pageNumber="61">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
as gracile are in
|
||
<emphasis box="[1182,1318,1104,1125]" italics="true" pageId="60" pageNumber="61">boldface italics</emphasis>
|
||
with a boldface
|
||
<emphasis box="[1474,1483,1104,1126]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
G
|
||
<emphasis box="[1500,1509,1104,1126]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
; specimens considered in
|
||
<bibRefCitation author="Larson P." box="[769,903,1133,1154]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[769,903,1133,1154]" italics="true" pageId="60" pageNumber="61">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
as robust are in boldface with a boldface
|
||
<emphasis box="[1318,1327,1133,1155]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
R
|
||
<emphasis box="[1342,1351,1133,1155]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
; specimens considered in
|
||
<bibRefCitation author="Larson P." box="[626,759,1162,1184]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[626,759,1162,1184]" italics="true" pageId="60" pageNumber="61">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
as referable to
|
||
<emphasis box="[908,932,1162,1184]" italics="true" pageId="60" pageNumber="61">“T</emphasis>
|
||
.
|
||
<emphasis box="[944,984,1162,1184]" italics="true" pageId="60" pageNumber="61">“x””</emphasis>
|
||
are in
|
||
<emphasis box="[1053,1107,1162,1183]" italics="true" pageId="60" pageNumber="61">italics</emphasis>
|
||
and marked with an
|
||
<emphasis box="[1318,1327,1162,1184]" italics="true" pageId="60" pageNumber="61">“</emphasis>
|
||
X
|
||
<emphasis box="[1343,1352,1162,1184]" italics="true" pageId="60" pageNumber="61">”</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="60.[524,1516,957,1213]" box="[1099,1515,1191,1213]" pageId="60" pageNumber="61">Full-size DOI: 10.7717/peerj.9192/fig-21</paragraph>
|
||
</caption>
|
||
<paragraph blockId="60.[498,1542,1268,1893]" pageId="60" pageNumber="61">
|
||
Several lines of evidence were offered to support the inference of a binary difference between male and female morphs (
|
||
<bibRefCitation author="Larson P." box="[914,1064,1667,1693]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="60" pageNumber="61" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[914,1064,1667,1693]" italics="true" pageId="60" pageNumber="61">Larson, 2008</emphasis>
|
||
</bibRefCitation>
|
||
). However, it was found here that those purported dimorphic features actually show a continuum of variation (e.g., circumference to length ratio of the humerus, height to length ratio of the ilium, circumference to length ratio of metatarsal II), whereas others show no pattern (e.g., ratio of third to second dentary teeth, divergence of the ischium, circumference to length ratio of the femur) (
|
||
<figureCitation box="[508,588,1866,1892]" captionStart="Figure 22" captionStartId="61.[524,589,955,977]" captionTargetBox="[491,1463,232,936]" captionTargetId="figure-236@61.[535,1005,247,893]" captionTargetPageId="61" captionText="Figure 22 Sex dimorphs of Larson (2008) mapped onto the growth curve of Tyrannosaurus rex. A transitional pattern is not seen between gracile and robust morphs; if sexual dimorphism was pre- sent, then the “gracile” and “robust” morphs should grade into each other, which is not seen. Likewise, an ontogenetic progression among the cranial and postcranial indices is not seen. See text for details. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-22" figureDoi="http://doi.org/10.5281/zenodo.5825016" httpUri="https://zenodo.org/record/5825016/files/figure.png" pageId="60" pageNumber="61">Fig. 22</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825016" ID-Zenodo-Dep="5825016" httpUri="https://zenodo.org/record/5825016/files/figure.png" pageId="61" pageNumber="62" startId="61.[524,589,955,977]" targetBox="[491,1463,232,936]" targetPageId="61">
|
||
<paragraph blockId="61.[524,1516,955,1094]" pageId="61" pageNumber="62">
|
||
<emphasis bold="true" box="[524,1515,955,977]" pageId="61" pageNumber="62">
|
||
Figure 22 Sex dimorphs of
|
||
<bibRefCitation author="Larson P." box="[812,953,955,976]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="61" pageNumber="62" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis bold="true" box="[812,953,955,976]" italics="true" pageId="61" pageNumber="62">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1320,1509,955,976]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="61" pageNumber="62" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1320,1509,955,976]" italics="true" pageId="61" pageNumber="62">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
A transitional pattern is not seen between gracile and robust morphs; if sexual dimorphism was present, then the
|
||
<emphasis box="[656,665,1013,1035]" italics="true" pageId="61" pageNumber="62">“</emphasis>
|
||
gracile
|
||
<emphasis box="[728,737,1013,1035]" italics="true" pageId="61" pageNumber="62">”</emphasis>
|
||
and
|
||
<emphasis box="[783,792,1013,1035]" italics="true" pageId="61" pageNumber="62">“</emphasis>
|
||
robust
|
||
<emphasis box="[854,863,1013,1035]" italics="true" pageId="61" pageNumber="62">”</emphasis>
|
||
morphs should grade into each other, which is not seen. Likewise, an ontogenetic progression among the cranial and postcranial indices is not seen. See text for details. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-22
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="61.[498,1542,1149,1654]" pageId="61" pageNumber="62">
|
||
<bibRefCitation author="Larson P." box="[530,691,1149,1175]" editor="Larson PL & Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="61" pageNumber="62" pagination="103 - 127" refId="ref54789" refString="Larson P. 2008. Variation and sexual dimorphism in Tyrannosaurus rex. In: Larson PL, Carpenter K, eds. Tyrannosaurus rex, the Tyrant King. Bloomington: Indiana University Press, 103 - 127." title="Variation and sexual dimorphism in Tyrannosaurus rex" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">
|
||
<emphasis box="[530,691,1149,1175]" italics="true" pageId="61" pageNumber="62">Larson (2008)</emphasis>
|
||
</bibRefCitation>
|
||
referred three specimens (
|
||
<materialsCitation box="[1002,1238,1149,1176]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="61" pageNumber="62" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1250,1367,1149,1175]" collectionCode="MOR" pageId="61" pageNumber="62" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1379,1534,1149,1175]" collectionCode="SDSM" pageId="61" pageNumber="62" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
) to the taxon
|
||
<emphasis box="[650,894,1189,1215]" italics="true" pageId="61" pageNumber="62">
|
||
“
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[662,837,1190,1215]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="61" pageNumber="62" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
“x””
|
||
</emphasis>
|
||
. If this taxon was valid, then the specimens should be recovered sharing a single branch, separate from the other adults. However, the specimens are scattered throughout the adult growth category of the ontogram, indicating that the characters thought to unite them (size of lacrimal pneumatic foramen, maxillary and dentary tooth count, form of second dentary tooth, ratio of the length of the third to second dentary tooth) actually reflect ontogenetic or individual variation, not taxonomically informative characters (
|
||
<figureCitation box="[953,1032,1428,1454]" captionStart="Figure 22" captionStartId="61.[524,589,955,977]" captionTargetBox="[491,1463,232,936]" captionTargetId="figure-236@61.[535,1005,247,893]" captionTargetPageId="61" captionText="Figure 22 Sex dimorphs of Larson (2008) mapped onto the growth curve of Tyrannosaurus rex. A transitional pattern is not seen between gracile and robust morphs; if sexual dimorphism was pre- sent, then the “gracile” and “robust” morphs should grade into each other, which is not seen. Likewise, an ontogenetic progression among the cranial and postcranial indices is not seen. See text for details. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-22" figureDoi="http://doi.org/10.5281/zenodo.5825016" httpUri="https://zenodo.org/record/5825016/files/figure.png" pageId="61" pageNumber="62">Fig. 22</figureCitation>
|
||
). Finally, a comparison of individual variation (Data S5) was done to identify any shared characters that might provide evidence of common identity. Individual variation was recovered for
|
||
<materialsCitation box="[1206,1439,1507,1534]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="61" pageNumber="62" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
and
|
||
<materialsCitation box="[498,654,1547,1573]" collectionCode="SDSM" pageId="61" pageNumber="62" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
, but not for
|
||
<materialsCitation box="[805,924,1547,1573]" collectionCode="MOR" pageId="61" pageNumber="62" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
. In addition to that, there are no shared characters between the two former specimens; therefore, no character evidence was found that supports the purported taxon.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="62" lastPageNumber="63" pageId="61" pageNumber="62" type="reference_group">
|
||
<paragraph blockId="61.[498,1542,1698,1890]" box="[498,730,1698,1728]" pageId="61" pageNumber="62">
|
||
<heading bold="true" box="[498,730,1698,1728]" fontSize="12" level="2" pageId="61" pageNumber="62" reason="0">
|
||
<emphasis bold="true" box="[498,730,1698,1728]" pageId="61" pageNumber="62">Dentary groove</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="61.[498,1542,1698,1890]" lastBlockId="62.[498,1542,234,898]" lastPageId="62" lastPageNumber="63" pageId="61" pageNumber="62">
|
||
It has been claimed by
|
||
<bibRefCitation author="Schmerge JD & Rothschild DM" box="[762,1120,1744,1770]" journalOrPublisher="Cretaceous Research" pageId="61" pageNumber="62" pagination="26 - 33" part="61" refId="ref55842" refString="Schmerge JD, Rothschild DM. 2016 a. Distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988. Cretaceous Research 61: 26 - 33 DOI 10.1016 / j. cretres. 2015.12.016." title="Distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988" type="journal article" year="2016">
|
||
<emphasis box="[762,1120,1744,1770]" italics="true" pageId="61" pageNumber="62">Schmerge & Rothschild (2016a)</emphasis>
|
||
</bibRefCitation>
|
||
that the neurovascular groove along the alveolar row of foramina of the dentary in tyrannosaurids has a binary expression of present and absent. Among latest Cretaceous tyrannosaurids of Laramidia, they hypothesized that the groove is absent from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1015,1082,1864,1889]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="61" pageNumber="62" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1015,1082,1864,1889]" italics="true" pageId="61" pageNumber="62">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, regardless of size, whereas it is present in
|
||
<emphasis box="[530,832,234,260]" italics="true" pageId="62" pageNumber="63">
|
||
“
|
||
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" box="[542,820,234,260]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="62" pageNumber="63" phylum="Chordata" rank="species" species="lancensis">Nanotyrannus lancensis</taxonomicName>
|
||
”
|
||
</emphasis>
|
||
, which otherwise has been shown to be an invalid taxon (
|
||
<bibRefCitation author="Carr TD & Williamson TE" box="[508,806,274,301]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="62" pageNumber="63" pagination="479 - 523" part="142" refId="ref52732" refString="Carr TD, Williamson TE. 2004. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142 (4): 479 - 523 DOI 10.1111 / j. 1096 - 3642.2004.00130. x." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">
|
||
<emphasis box="[508,806,274,301]" italics="true" pageId="62" pageNumber="63">Carr & Williamson, 2004</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="80" lastPageNumber="81" pageId="62" pageNumber="63" type="description">
|
||
<paragraph blockId="62.[498,1542,234,898]" pageId="62" pageNumber="63">
|
||
Several of the
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[698,767,315,340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="62" pageNumber="63" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[698,767,315,340]" italics="true" pageId="62" pageNumber="63">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens that
|
||
<bibRefCitation author="Schmerge JD & Rothschild DM" box="[959,1314,314,340]" journalOrPublisher="Cretaceous Research" pageId="62" pageNumber="63" pagination="26 - 33" part="61" refId="ref55842" refString="Schmerge JD, Rothschild DM. 2016 a. Distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988. Cretaceous Research 61: 26 - 33 DOI 10.1016 / j. cretres. 2015.12.016." title="Distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988" type="journal article" year="2016">
|
||
<emphasis box="[959,1314,314,340]" italics="true" pageId="62" pageNumber="63">Schmerge & Rothschild (2016a</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<emphasis box="[1329,1408,314,340]" italics="true" pageId="62" pageNumber="63">
|
||
<bibRefCitation author="Schmerge JD & Rothschild DM" box="[1329,1400,314,340]" journalOrPublisher="Cretaceous Research" pageId="62" pageNumber="63" pagination="238 - 243" part="65" refId="ref55893" refString="Schmerge JD, Rothschild DM. 2016 b. When a groove is not a groove: clarification of the appearance of the dentary groove in tyrannosauroid theropods and the distinction between Nanotyrannus and Tyrannosaurus. Reply to comment on: distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988. Cretaceous Research 65: 238 - 243 DOI 10.1016 / j. cretres. 2016.04.015." title="When a groove is not a groove: clarification of the appearance of the dentary groove in tyrannosauroid theropods and the distinction between Nanotyrannus and Tyrannosaurus. Reply to comment on: distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988" type="journal article" year="2016">2016b</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
claimed to lack the dentary groove were re-examined for this study and it was found that in two of them the groove is present, but greatly reduced in extent (
|
||
<materialsCitation box="[1166,1400,393,420]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="62" pageNumber="63" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1411,1529,394,420]" collectionCode="MOR" pageId="62" pageNumber="63" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
). In addition to that, a reduced groove is seen in all adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1138,1204,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="62" pageNumber="63" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1138,1204,434,459]" italics="true" pageId="62" pageNumber="63">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, including the type specimen (
|
||
<materialsCitation box="[506,620,473,499]" collectionCode="CM" pageId="62" pageNumber="63" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
,
|
||
<materialsCitation box="[633,795,473,499]" collectionCode="LACM" pageId="62" pageNumber="63" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<materialsCitation box="[808,926,473,500]" collectionCode="MOR" pageId="62" pageNumber="63" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
,
|
||
<materialsCitation box="[939,1057,473,499]" collectionCode="MOR" pageId="62" pageNumber="63" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1070,1202,473,500]" collectionCode="MOR" pageId="62" pageNumber="63" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1215,1388,473,500]" collectionCode="UWBM" pageId="62" pageNumber="63" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
) (Data S1). These grooves are true sulci, and are not artifacts of foramen shape, and so are not
|
||
<emphasis box="[498,510,553,579]" italics="true" pageId="62" pageNumber="63">“</emphasis>
|
||
pseudo-grooves
|
||
<emphasis box="[693,705,553,579]" italics="true" pageId="62" pageNumber="63">”</emphasis>
|
||
(sensu
|
||
<bibRefCitation author="Schmerge JD & Rothschild DM" box="[796,1148,553,579]" journalOrPublisher="Cretaceous Research" pageId="62" pageNumber="63" pagination="238 - 243" part="65" refId="ref55893" refString="Schmerge JD, Rothschild DM. 2016 b. When a groove is not a groove: clarification of the appearance of the dentary groove in tyrannosauroid theropods and the distinction between Nanotyrannus and Tyrannosaurus. Reply to comment on: distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988. Cretaceous Research 65: 238 - 243 DOI 10.1016 / j. cretres. 2016.04.015." title="When a groove is not a groove: clarification of the appearance of the dentary groove in tyrannosauroid theropods and the distinction between Nanotyrannus and Tyrannosaurus. Reply to comment on: distribution of the dentary groove of theropod dinosaurs: implications for theropod phylogeny and the validity of the genus Nanotyrannus Bakker et al. 1988" type="journal article" year="2016">
|
||
<emphasis box="[796,1148,553,579]" italics="true" pageId="62" pageNumber="63">Schmerge & Rothschild, 2016b</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1542,234,898]" pageId="62" pageNumber="63">
|
||
Although the states of this character were not unambiguously optimized on the ontogram, the specimens show a stepwise progression: the groove is absent from the least mature specimen (
|
||
<materialsCitation box="[716,882,673,699]" collectionCode="LACM" pageId="62" pageNumber="63" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
), the groove is present along the entire row in more mature juveniles (e.g., BMRP 2002.4.1), and, finally, the groove is reduced to short segments in adults (e.g.,
|
||
<materialsCitation box="[789,975,752,779]" collectionCode="FMNH" pageId="62" pageNumber="63" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
) and so cannot be said to be absent from them. The groove is almost certainly obscured in adults by the overall expansion of the bone in response to growth, higher loads imposed by ontogenetic increase in body size, tooth size, and bite force (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[737,1046,872,898]" journalOrPublisher="Biology Letters" pageId="62" pageNumber="63" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[737,1046,872,898]" italics="true" pageId="62" pageNumber="63">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1542,938,1249]" box="[498,1254,938,968]" pageId="62" pageNumber="63">
|
||
<heading bold="true" box="[498,1254,938,968]" fontSize="12" level="2" pageId="62" pageNumber="63" reason="0">
|
||
<emphasis bold="true" box="[498,1254,938,968]" pageId="62" pageNumber="63">Synthesis of ontogeny and functional morphology</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1542,938,1249]" pageId="62" pageNumber="63">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,718,984,1009]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="62" pageNumber="63" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,718,984,1009]" italics="true" pageId="62" pageNumber="63">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
has been central in recent quantitative studies of the functional morphology of tyrannosaurids and other large theropods (
|
||
<bibRefCitation author="Henderson DM" box="[1180,1374,1023,1049]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="62" pageNumber="63" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[1180,1374,1023,1049]" italics="true" pageId="62" pageNumber="63">Henderson, 2002</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Henderson DM & Snively E." journalOrPublisher="Proceedings of the Royal Society B" pageId="62" pageNumber="63" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="62" pageNumber="63">Henderson & Snively, 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[665,1063,1063,1090]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="62" pageNumber="63" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis box="[665,1063,1063,1090]" italics="true" pageId="62" pageNumber="63">Therrien, Henderson & Ruff, 2005</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1078,1493,1063,1090]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="62" pageNumber="63" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1078,1493,1063,1090]" italics="true" pageId="62" pageNumber="63">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[498,710,1103,1129]" journalOrPublisher="PeerJ" pageId="62" pageNumber="63" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[498,710,1103,1129]" italics="true" pageId="62" pageNumber="63">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
). Those results are re-examined here in the framework of the ontogram to assess the completeness of the hypotheses, provide a specific ontogenetic context for major functional changes, identify the gaps in the data, and propose hypotheses of where in the growth series major morphological changes occurred.
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1542,1282,1791]" box="[498,971,1282,1310]" pageId="62" pageNumber="63">
|
||
<heading bold="true" box="[498,971,1282,1310]" fontSize="11" level="3" pageId="62" pageNumber="63" reason="6">
|
||
<emphasis bold="true" box="[498,971,1282,1310]" italics="true" pageId="62" pageNumber="63">
|
||
Orbital fenestra (
|
||
<bibRefCitation author="Henderson DM" box="[730,962,1282,1310]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="62" pageNumber="63" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">Henderson, 2002</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1542,1282,1791]" pageId="62" pageNumber="63">
|
||
Quantifiable correlates of the height of the skull frame include the size, shape, and orientation of the orbital fenestra, the width of the pre- and postorbital bars that frame it, and the stress regime of the skull at the orbit (
|
||
<bibRefCitation author="Henderson DM" box="[1048,1243,1406,1432]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="62" pageNumber="63" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[1048,1243,1406,1432]" italics="true" pageId="62" pageNumber="63">Henderson, 2002</emphasis>
|
||
</bibRefCitation>
|
||
). One juvenile (
|
||
<materialsCitation collectionCode="CMNH" pageId="62" pageNumber="63" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
) and adult (
|
||
<materialsCitation box="[702,939,1446,1473]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="62" pageNumber="63" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
) were compared by
|
||
<bibRefCitation author="Henderson DM" box="[1181,1378,1446,1472]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="62" pageNumber="63" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[1181,1378,1446,1472]" italics="true" pageId="62" pageNumber="63">Henderson (2002</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1394,1473,1446,1472]" captionStart="Figure 23" captionStartId="63.[524,589,955,977]" captionTargetBox="[485,1471,224,936]" captionTargetId="figure-262@63.[535,1005,247,893]" captionTargetPageId="63" captionText="Figure 23 The results of Henderson (2002) mapped onto the growth curve of Tyrannosaurus rex. All measures of the correlates of orbital fenestra size and shape change from juvenile to adult categories. It is predicted here that this transition occurred early in ontogeny, at the subadult growth stage, given the presence of correlates of a tall skull in subadult specimens. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-23" figureDoi="http://doi.org/10.5281/zenodo.5825020" httpUri="https://zenodo.org/record/5825020/files/figure.png" pageId="62" pageNumber="63">Fig. 23</figureCitation>
|
||
). Comparison of the results of
|
||
<bibRefCitation author="Henderson DM" box="[845,1052,1485,1512]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="62" pageNumber="63" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[845,1052,1485,1512]" italics="true" pageId="62" pageNumber="63">Henderson (2002)</emphasis>
|
||
</bibRefCitation>
|
||
with the growth curve reconstructed here (
|
||
<figureCitation box="[566,644,1525,1552]" captionStart="Figure 23" captionStartId="63.[524,589,955,977]" captionTargetBox="[485,1471,224,936]" captionTargetId="figure-262@63.[535,1005,247,893]" captionTargetPageId="63" captionText="Figure 23 The results of Henderson (2002) mapped onto the growth curve of Tyrannosaurus rex. All measures of the correlates of orbital fenestra size and shape change from juvenile to adult categories. It is predicted here that this transition occurred early in ontogeny, at the subadult growth stage, given the presence of correlates of a tall skull in subadult specimens. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-23" figureDoi="http://doi.org/10.5281/zenodo.5825020" httpUri="https://zenodo.org/record/5825020/files/figure.png" pageId="62" pageNumber="63">Fig. 23</figureCitation>
|
||
) shows that the details of the transition are incomplete, including the precise stage at which the transformation occurred owing to the absence of suitably complete subadult specimens from the current sample in museum collections. Given the presence of correlates of a tall skull in the exemplars of the subadult category (
|
||
<materialsCitation box="[1285,1449,1645,1672]" collectionCode="LACM" pageId="62" pageNumber="63" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="RSM" pageId="62" pageNumber="63" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
) it is predicted here that the reduction in orbit size, increase in ellipticity, increase in vertical orientation, increase in pre- and postorbital bar length, and increase in stress regime at the orbit first occurred abruptly in subadults.
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1541,1824,1935]" box="[498,1292,1824,1852]" pageId="62" pageNumber="63">
|
||
<emphasis bold="true" box="[498,1292,1824,1852]" italics="true" pageId="62" pageNumber="63">
|
||
Skull & jaw strength (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[794,1282,1824,1852]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="62" pageNumber="63" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">Snively, Henderson & Phillips, 2006</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="62.[498,1541,1824,1935]" lastBlockId="63.[498,1542,1149,1893]" lastPageId="63" lastPageNumber="64" pageId="62" pageNumber="63">
|
||
A study of the bending strength of the skull and teeth of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1175,1244,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="62" pageNumber="63" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1175,1244,1869,1894]" italics="true" pageId="62" pageNumber="63">T. rex</emphasis>
|
||
</taxonomicName>
|
||
by
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" journalOrPublisher="Acta Palaeontologica Polonica" pageId="62" pageNumber="63" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="62" pageNumber="63">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
included comparison between one juvenile (BMRP 2002.4.1), two adults (
|
||
<materialsCitation box="[506,744,1149,1176]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="63" pageNumber="64" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[757,920,1149,1175]" collectionCode="LACM" pageId="63" pageNumber="64" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
), and one senescent adult (
|
||
<materialsCitation box="[1240,1432,1149,1175]" collectionCode="FMNH" pageId="63" pageNumber="64" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
). Their results were mapped onto the growth curve to obtain an ontogenetically constrained hypothesis of the comparisons of vertical bending strength, lateral bending strength, torsional strength, and tooth bending strength (
|
||
<figureCitation box="[1062,1142,1268,1295]" captionStart="Figure 24" captionStartId="64.[524,589,954,976]" captionTargetBox="[483,1457,232,934]" captionTargetId="figure-273@64.[536,1005,247,892]" captionTargetPageId="64" captionText="Figure 24 Skull bending strength mapped onto the growth curve of Tyrannosaurus rex. The results of Snively, Henderson & Phillips (2006) showing that the subadult growth stage was an important functional transition point during ontogeny between the long and low skulls of adults and tall and sturdy skulls of more mature animals. Their results show a progression in strength of the skull frame and dentition throughout the adult categories. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-24" figureDoi="http://doi.org/10.5281/zenodo.5825022" httpUri="https://zenodo.org/record/5825022/files/figure.png" pageId="63" pageNumber="64">Fig. 24</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825020" ID-Zenodo-Dep="5825020" httpUri="https://zenodo.org/record/5825020/files/figure.png" pageId="63" pageNumber="64" startId="63.[524,589,955,977]" targetBox="[485,1471,224,936]" targetPageId="63">
|
||
<paragraph blockId="63.[524,1515,954,1094]" pageId="63" pageNumber="64">
|
||
<emphasis bold="true" box="[524,1481,954,977]" pageId="63" pageNumber="64">
|
||
Figure 23 The results of
|
||
<bibRefCitation author="Henderson DM" box="[771,947,954,976]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="63" pageNumber="64" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis bold="true" box="[771,947,954,976]" italics="true" pageId="63" pageNumber="64">Henderson (2002)</emphasis>
|
||
</bibRefCitation>
|
||
mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1290,1475,955,976]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="63" pageNumber="64" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1290,1475,955,976]" italics="true" pageId="63" pageNumber="64">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
All measures of the correlates of orbital fenestra size and shape change from juvenile to adult categories. It is predicted here that this transition occurred early in ontogeny, at the subadult growth stage, given the presence of correlates of a tall skull in subadult specimens. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-23
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="63.[498,1542,1149,1893]" pageId="63" pageNumber="64">
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[530,958,1308,1335]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="63" pageNumber="64" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[530,958,1308,1335]" italics="true" pageId="63" pageNumber="64">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
provided a summary hypothesis of the phylogenetic progression of skull strengthening attributes of tyrannosaurids. This phylogenetic progression was compared with the ontogenetic progression for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1398,1466,1389,1414]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="63" pageNumber="64" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1398,1466,1389,1414]" italics="true" pageId="63" pageNumber="64">T. rex</emphasis>
|
||
</taxonomicName>
|
||
found here to test the hypothesis of congruence between ontogeny and phylogeny. The sequences of changes are congruent, aside from the occurrence of a wide adductor chamber; in
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[498,926,1507,1534]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="63" pageNumber="64" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[498,926,1507,1534]" italics="true" pageId="63" pageNumber="64">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
this character is unique to
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1251,1320,1508,1534]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="63" pageNumber="64" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1251,1268,1508,1533]" italics="true" pageId="63" pageNumber="64">T</emphasis>
|
||
.
|
||
<emphasis box="[1284,1320,1508,1533]" italics="true" pageId="63" pageNumber="64">rex</emphasis>
|
||
</taxonomicName>
|
||
, and is the last character to evolve, whereas it is present in the second growth stage as shown by its presence in
|
||
<materialsCitation box="[640,796,1587,1614]" collectionCode="CMNH" pageId="63" pageNumber="64" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
, a juvenile, and all specimens that are more mature than it.
|
||
</paragraph>
|
||
<paragraph blockId="63.[498,1542,1149,1893]" lastBlockId="64.[498,1542,1177,1921]" lastPageId="64" lastPageNumber="65" pageId="63" pageNumber="64">
|
||
As in the phylogenetic transition, fused nasals are seen earliest, rostrally wide nasals are seen in large juveniles, and the increased nasal cross section, peg-in-socket nasomaxillary suture, and increased tooth strength occur simultaneously in subadults. The simultaneity is almost certainly an artifact of a low sample size, and a higher sample of subadults is required to test the hypothesis that acquisition of those features occur in the same order as is seen in the phylogenetic sequence. All of these features occur in the growth of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[621,690,1867,1892]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="63" pageNumber="64" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[621,690,1867,1892]" italics="true" pageId="63" pageNumber="64">T. rex</emphasis>
|
||
</taxonomicName>
|
||
before the 3,000 kg threshold is reached, indicating that they are not an artifact of the descendant (i.e.,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[864,932,1178,1203]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[864,932,1178,1203]" italics="true" pageId="64" pageNumber="65">T. rex</emphasis>
|
||
</taxonomicName>
|
||
) size range; indeed, these features are seen in other, smaller tyrannosaurines (e.g.,
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[928,1103,1217,1243]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="genus">
|
||
<emphasis box="[928,1103,1217,1243]" italics="true" pageId="64" pageNumber="65">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority=", Zhuchengtyrannus" authorityName="Zhuchengtyrannus" box="[1118,1457,1217,1244]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1118,1135,1218,1243]" italics="true" pageId="64" pageNumber="65">T</emphasis>
|
||
.
|
||
<emphasis box="[1150,1226,1217,1243]" italics="true" pageId="64" pageNumber="65">bataar</emphasis>
|
||
,
|
||
<emphasis box="[1242,1457,1217,1243]" italics="true" pageId="64" pageNumber="65">Zhuchengtyrannus</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825022" ID-Zenodo-Dep="5825022" httpUri="https://zenodo.org/record/5825022/files/figure.png" pageId="64" pageNumber="65" startId="64.[524,589,954,976]" targetBox="[483,1457,232,934]" targetPageId="64">
|
||
<paragraph blockId="64.[524,1515,954,1122]" pageId="64" pageNumber="65">
|
||
<emphasis bold="true" box="[524,1382,954,976]" pageId="64" pageNumber="65">
|
||
Figure 24 Skull bending strength mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1191,1376,954,975]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1191,1376,954,975]" italics="true" pageId="64" pageNumber="65">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The results of
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[524,866,983,1005]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="64" pageNumber="65" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[524,866,983,1005]" italics="true" pageId="64" pageNumber="65">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
showing that the subadult growth stage was an important functional transition point during ontogeny between the long and low skulls of adults and tall and sturdy skulls of more mature animals. Their results show a progression in strength of the skull frame and dentition throughout the adult categories. Key to specimens numbered on the growth curve is in Fig. 12.
|
||
</paragraph>
|
||
<paragraph blockId="64.[524,1515,954,1122]" box="[1099,1515,1100,1122]" pageId="64" pageNumber="65">Full-size DOI: 10.7717/peerj.9192/fig-24</paragraph>
|
||
</caption>
|
||
<paragraph blockId="64.[498,1542,1177,1921]" pageId="64" pageNumber="65">
|
||
The data for maxillary tooth bending strength are from the juvenile (BMRP 2002.4.1) and adult (
|
||
<materialsCitation box="[621,854,1297,1324]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="64" pageNumber="65" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[864,1022,1297,1323]" collectionCode="LACM" pageId="64" pageNumber="65" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
) stages (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1121,1526,1297,1323]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="64" pageNumber="65" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1121,1526,1297,1323]" italics="true" pageId="64" pageNumber="65">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
), which corresponds to the 5th, 14th, and 18th growth stages of the ontogram (
|
||
<figureCitation box="[508,584,1376,1403]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="64" pageNumber="65">Figs. 2</figureCitation>
|
||
and 24). Labiolingual tooth bending strength increases by an order of magnitude between the juvenile and adult growth stages; the juvenile bending strength is comparable to those of juvenile
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[733,998,1456,1482]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[733,998,1456,1482]" italics="true" pageId="64" pageNumber="65">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="(Snively, Henderson & Phillips, 2006)" baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1059,1316,1456,1482]" italics="true" pageId="64" pageNumber="65">Albertosaurus libratus</emphasis>
|
||
(
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" journalOrPublisher="Acta Palaeontologica Polonica" pageId="64" pageNumber="65" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="64" pageNumber="65">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, which are lower than those of nontyrannosaurids.
|
||
</paragraph>
|
||
<paragraph blockId="64.[498,1542,1177,1921]" lastBlockId="65.[498,1542,234,1456]" lastPageId="65" lastPageNumber="66" pageId="64" pageNumber="65">
|
||
Mesiodistal bending strength increases from juvenile to adult, where that of the juvenile is most comparable to narrow-toothed
|
||
<taxonomicName authority=", Monolophosaurus" authorityName="Monolophosaurus" box="[962,1306,1576,1602]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="genus">
|
||
<emphasis box="[962,1084,1576,1602]" italics="true" pageId="64" pageNumber="65">Allosaurus</emphasis>
|
||
,
|
||
<emphasis box="[1099,1306,1576,1602]" italics="true" pageId="64" pageNumber="65">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Snively, Henderson & Phillips, 2006)" baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="64" pageNumber="65" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis italics="true" pageId="64" pageNumber="65">Albertosaurus libratus</emphasis>
|
||
(
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[605,1020,1615,1642]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="64" pageNumber="65" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[605,1020,1615,1642]" italics="true" pageId="64" pageNumber="65">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. This transition in tooth strength almost certainly occurred between the 5th and 6th growth stages since the 7-shaped lacrimal (
|
||
<materialsCitation box="[506,656,1695,1721]" collectionCode="RSM" pageId="64" pageNumber="65" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
), regarded here as a correlate of high bites forces, is seen at the 6th growth stage, and wide teeth (
|
||
<materialsCitation box="[840,1001,1735,1762]" collectionCode="LACM" pageId="64" pageNumber="65" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
) are present no later than growth stage 7. Also, the correlates of high bite forces identified by
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1042,1470,1775,1801]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="64" pageNumber="65" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1042,1470,1775,1801]" italics="true" pageId="64" pageNumber="65">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
, including a peg-in-socket nasomaxillary suture, is present in subadults (
|
||
<materialsCitation box="[1323,1486,1815,1842]" collectionCode="LACM" pageId="64" pageNumber="65" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
); the other correlates, such as a tall maxilla with a wide palatal shelf are unequivocally present by growth stage 8 (
|
||
<materialsCitation box="[687,821,1894,1921]" collectionCode="MOR" pageId="64" pageNumber="65" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
) and are predicted here to be present in subadults once more complete specimens are found. If true, then the teeth, along with the entire craniodental mechanism rapidly (<3 years) transformed as an integrated system well in advance of somatic maturity. The increase in both tooth strength indicators among adults indicates that this trend of increasing strength continued into later growth stages; however, this observation is based on two specimens (
|
||
<materialsCitation box="[1127,1362,393,420]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="65" pageNumber="66" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1374,1534,394,420]" collectionCode="LACM" pageId="65" pageNumber="66" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
) and data from additional adults are required to test that hypothesis (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,234,1456]" pageId="65" pageNumber="66">
|
||
Juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[631,699,514,539]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="65" pageNumber="66" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[631,699,514,539]" italics="true" pageId="65" pageNumber="66">T. rex</emphasis>
|
||
</taxonomicName>
|
||
started life with snouts that are strengthened by fusion of the internasal suture and the vaulted cross section of the nasals (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1095,1510,553,579]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1095,1510,553,579]" italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
). However, the dorsoventrally shallow skull and the wide and flat frontal ramus of the nasals produces a low vertical bending strength (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1075,1490,633,659]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1075,1490,633,659]" italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
). The tongue-in-groove nasomaxillary joint surface and the narrow span across the nasals are correlates of a low lateral bending and low torsional strength (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,234,1456]" pageId="65" pageNumber="66">
|
||
Based on their observations of adults,
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[981,1409,792,819]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[981,1409,792,819]" italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
identified two correlates of high torsional strength and lateral bending strength of the skull: a peg-insocket nasomaxillary suture and a wide span across rostral end of the nasals, which are seen in the subadult category (
|
||
<materialsCitation box="[894,1058,911,938]" collectionCode="LACM" pageId="65" pageNumber="66" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
), and so the transition from a weak snout to a strong snout occurred in that earlier growth stage and was carried forward into adulthood. The change from a low to a tall skull is a central part of the transition from a weak to a strong skull (
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[770,1181,1031,1058]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[770,1181,1031,1058]" italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
); correlates of a tall skull, such as a wide preorbital bar (
|
||
<materialsCitation box="[793,941,1071,1097]" collectionCode="RSM" pageId="65" pageNumber="66" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
) in the subadult growth stage, shows that the transition from shallow to tall also occurred from the juvenile to the subadult growth categories.
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,234,1456]" pageId="65" pageNumber="66">
|
||
As seen in the maxillary teeth, a trend of increased bending strength of the skull is seen among adults when the results of
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[882,1298,1230,1257]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="65" pageNumber="66" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[882,1298,1230,1257]" italics="true" pageId="65" pageNumber="66">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
are mapped onto the ontogram (
|
||
<figureCitation box="[631,711,1270,1296]" captionStart="Figure 24" captionStartId="64.[524,589,954,976]" captionTargetBox="[483,1457,232,934]" captionTargetId="figure-273@64.[536,1005,247,892]" captionTargetPageId="64" captionText="Figure 24 Skull bending strength mapped onto the growth curve of Tyrannosaurus rex. The results of Snively, Henderson & Phillips (2006) showing that the subadult growth stage was an important functional transition point during ontogeny between the long and low skulls of adults and tall and sturdy skulls of more mature animals. Their results show a progression in strength of the skull frame and dentition throughout the adult categories. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-24" figureDoi="http://doi.org/10.5281/zenodo.5825022" httpUri="https://zenodo.org/record/5825022/files/figure.png" pageId="65" pageNumber="66">Fig. 24</figureCitation>
|
||
). However, only two specimens are sampled (
|
||
<materialsCitation box="[1255,1492,1270,1297]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="65" pageNumber="66" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,683,1310,1337]" collectionCode="FMNH" pageId="65" pageNumber="66" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
) and so additional adult specimens, and juveniles, are required to test this hypothesis. It is predicted that the trend of increasing bending strengths occurred throughout ontogeny, with an abrupt increase at the transition between juveniles and subadults.
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,1495,1924]" box="[498,1359,1495,1523]" pageId="65" pageNumber="66">
|
||
<emphasis bold="true" box="[498,1359,1495,1523]" italics="true" pageId="65" pageNumber="66">
|
||
Mandibular ramus strength (
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[890,1350,1495,1523]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="65" pageNumber="66" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">Therrien, Henderson & Ruff, 2005</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,1495,1924]" pageId="65" pageNumber="66">
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[498,900,1539,1565]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="65" pageNumber="66" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis box="[498,900,1539,1565]" italics="true" pageId="65" pageNumber="66">Therrien, Henderson & Ruff (2005)</emphasis>
|
||
</bibRefCitation>
|
||
found that during the ontogeny of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1307,1374,1539,1564]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="65" pageNumber="66" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1307,1374,1539,1564]" italics="true" pageId="65" pageNumber="66">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, from juvenile to adult, the rostral end of the mandible could resist high torsional loads and, between growth stages, an increase in bite force was seen. Measurements were obtained from the mid-dentary and at the second alveolus (
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[1028,1426,1658,1685]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="65" pageNumber="66" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis box="[1028,1426,1658,1685]" italics="true" pageId="65" pageNumber="66">Therrien, Henderson & Ruff, 2005</emphasis>
|
||
</bibRefCitation>
|
||
). Two sets of measurements were taken, dorsoventral bending strength and relative strength (the ratio of dorsoventral and mediolateral bending strengths). The results of
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="65" pageNumber="66" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis italics="true" pageId="65" pageNumber="66">Therrien, Henderson & Ruff (2005)</emphasis>
|
||
</bibRefCitation>
|
||
were mapped onto the growth curve to test their hypotheses of ontogenetic change in the mandibular bending strengths of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1206,1274,1818,1843]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="65" pageNumber="66" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1206,1274,1818,1843]" italics="true" pageId="65" pageNumber="66">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[1293,1372,1817,1844]" captionStart="Figure 25" captionStartId="66.[524,589,953,975]" captionTargetBox="[483,1463,226,940]" captionTargetId="figure-289@66.[537,1006,247,892]" captionTargetPageId="66" captionText="Figure 25 The results of Therrien, Henderson & Ruff (2005) compared with the growth curve of Tyrannosaurus rex. Vertical bending strength and relative bending strength (sensu Therrien, Henderson & Ruff, 2005) mapped onto the growth curve of T. rex. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25" figureDoi="http://doi.org/10.5281/zenodo.5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="65" pageNumber="66">Fig. 25</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="65.[498,1542,1495,1924]" lastBlockId="66.[498,1542,1177,1921]" lastPageId="66" lastPageNumber="67" pageId="65" pageNumber="66">
|
||
In comparison with the growth curve, dorsoventral bending strength at the second alveolus shows a progressive increase from juveniles to young adults to adults, but the trend is lost among adults (
|
||
<figureCitation box="[814,891,1177,1204]" captionStart="Figure 25" captionStartId="66.[524,589,953,975]" captionTargetBox="[483,1463,226,940]" captionTargetId="figure-289@66.[537,1006,247,892]" captionTargetPageId="66" captionText="Figure 25 The results of Therrien, Henderson & Ruff (2005) compared with the growth curve of Tyrannosaurus rex. Vertical bending strength and relative bending strength (sensu Therrien, Henderson & Ruff, 2005) mapped onto the growth curve of T. rex. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25" figureDoi="http://doi.org/10.5281/zenodo.5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="66" pageNumber="67">Fig. 25</figureCitation>
|
||
). This loss of pattern is not unexpected for specimens of the variable adult size. It is predicted here that the strength of subadults will be lower than adults but will found to be closer to the adult values than to those of juveniles.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825024" ID-Zenodo-Dep="5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="66" pageNumber="67" startId="66.[524,589,953,975]" targetBox="[483,1463,226,940]" targetPageId="66">
|
||
<paragraph blockId="66.[524,1515,953,1122]" pageId="66" pageNumber="67">
|
||
<emphasis bold="true" pageId="66" pageNumber="67">
|
||
Figure 25 The results of
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[784,1142,953,975]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="66" pageNumber="67" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis bold="true" box="[784,1142,953,975]" italics="true" pageId="66" pageNumber="67">Therrien, Henderson & Ruff (2005)</emphasis>
|
||
</bibRefCitation>
|
||
compared with the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,720,983,1004]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="66" pageNumber="67" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[524,720,983,1004]" italics="true" pageId="66" pageNumber="67">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Vertical bending strength and relative bending strength (sensu
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="66" pageNumber="67" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis italics="true" pageId="66" pageNumber="67">Therrien, Henderson & Ruff, 2005</emphasis>
|
||
</bibRefCitation>
|
||
) mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1105,1161,1013,1034]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="66" pageNumber="67" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1105,1161,1013,1034]" italics="true" pageId="66" pageNumber="67">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="66.[498,1542,1177,1921]" pageId="66" pageNumber="67">
|
||
Although mid-dentary vertical bending strength is not available for juveniles, an increasing trend of strength is seen from the young adult to adult to senescent growth stages; in contrast, a clear pattern of increase is not seen among adults although the highest value is seen in the senescent adult (
|
||
<figureCitation box="[1016,1095,1416,1443]" captionStart="Figure 25" captionStartId="66.[524,589,953,975]" captionTargetBox="[483,1463,226,940]" captionTargetId="figure-289@66.[537,1006,247,892]" captionTargetPageId="66" captionText="Figure 25 The results of Therrien, Henderson & Ruff (2005) compared with the growth curve of Tyrannosaurus rex. Vertical bending strength and relative bending strength (sensu Therrien, Henderson & Ruff, 2005) mapped onto the growth curve of T. rex. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25" figureDoi="http://doi.org/10.5281/zenodo.5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="66" pageNumber="67">Fig. 25</figureCitation>
|
||
). It is predicted here that the bending strength of subadults at the mid-dentary will be less than those of the adult growth categories and that of juveniles will be significantly less than subadults.
|
||
</paragraph>
|
||
<paragraph blockId="66.[498,1542,1177,1921]" pageId="66" pageNumber="67">
|
||
Relative strength was also estimated at the second alveolus and at the mid-length of the dentary tooth row (
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[731,1129,1575,1602]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="66" pageNumber="67" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis box="[731,1129,1575,1602]" italics="true" pageId="66" pageNumber="67">Therrien, Henderson & Ruff, 2005</emphasis>
|
||
</bibRefCitation>
|
||
). Relative strength at the second alveolus increases from the juvenile to young adult growth categories, but a trend is not seen throughout the growth series (
|
||
<figureCitation box="[956,1034,1655,1682]" captionStart="Figure 25" captionStartId="66.[524,589,953,975]" captionTargetBox="[483,1463,226,940]" captionTargetId="figure-289@66.[537,1006,247,892]" captionTargetPageId="66" captionText="Figure 25 The results of Therrien, Henderson & Ruff (2005) compared with the growth curve of Tyrannosaurus rex. Vertical bending strength and relative bending strength (sensu Therrien, Henderson & Ruff, 2005) mapped onto the growth curve of T. rex. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25" figureDoi="http://doi.org/10.5281/zenodo.5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="66" pageNumber="67">Fig. 25</figureCitation>
|
||
). In addition to that, values for some adults are lower (e.g., 0.72) than that for the juvenile growth stage (e.g., 0.86), indicating a high degree of variation. It is predicted here that, if there is a trend, the relative strength of subadults will be intermediate between the values for the juvenile and young adult categories.
|
||
</paragraph>
|
||
<paragraph blockId="66.[498,1542,1177,1921]" lastBlockId="67.[498,1542,234,580]" lastPageId="67" lastPageNumber="68" pageId="66" pageNumber="67">
|
||
At the mid-dentary, relative strength decreases from young adults to adults, but no trend is seen among adults and senescent adults. It is predicted here, based on the trend in
|
||
<figureCitation box="[498,578,234,261]" captionStart="Figure 25" captionStartId="66.[524,589,953,975]" captionTargetBox="[483,1463,226,940]" captionTargetId="figure-289@66.[537,1006,247,892]" captionTargetPageId="66" captionText="Figure 25 The results of Therrien, Henderson & Ruff (2005) compared with the growth curve of Tyrannosaurus rex. Vertical bending strength and relative bending strength (sensu Therrien, Henderson & Ruff, 2005) mapped onto the growth curve of T. rex. Values for juveniles are missing for mid-dentary dorsoventral strength and mid-dentary relative strength. In general, strength increases ontogenetically, a trend that becomes obscured in adulthood. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-25" figureDoi="http://doi.org/10.5281/zenodo.5825024" httpUri="https://zenodo.org/record/5825024/files/figure.png" pageId="67" pageNumber="68">Fig. 25</figureCitation>
|
||
and the values for small juvenile tyrannosaurids in
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="67" pageNumber="68" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis italics="true" pageId="67" pageNumber="68">Therrien, Henderson & Ruff (2005)</emphasis>
|
||
</bibRefCitation>
|
||
, that the values for subadults and juveniles will be sequentially greater than those of the adult category.
|
||
</paragraph>
|
||
<paragraph blockId="67.[498,1542,234,580]" pageId="67" pageNumber="68">In short, vertical bending strength of the mandibular ramus increases ontogenetically, a trend that becomes obscured at senescence. In contrast, a trend in relative strength has a broadly decreasing trend at mid-dentary, but no clear trend at the second alveolus; that is, more variation is seen in relative strength than in dorsoventral strength. It is predicted that data from the future discovery of subadults will be consistent with the mid-dentary pattern but will not clarify the situation at the second alveolus.</paragraph>
|
||
<paragraph blockId="67.[498,1542,618,1525]" box="[498,1041,618,646]" pageId="67" pageNumber="68">
|
||
<heading bold="true" box="[498,1041,618,646]" fontSize="11" level="3" pageId="67" pageNumber="68" reason="6">
|
||
<emphasis bold="true" box="[498,1041,618,646]" italics="true" pageId="67" pageNumber="68">Summary of craniomandibular strength</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="67.[498,1542,618,1525]" pageId="67" pageNumber="68">
|
||
Taken together, the results of
|
||
<bibRefCitation author="Henderson DM" box="[844,1050,662,688]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="67" pageNumber="68" pagination="766 - 778" part="22" refId="ref54146" refString="Henderson DM. 2002. The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force. Journal of Vertebrate Paleontology 22 (4): 766 - 778 DOI 10.1671 / 0272 - 4634 (2002) 022 [0766: TEHITS] 2.0. CO; 2." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">
|
||
<emphasis box="[844,1050,662,688]" italics="true" pageId="67" pageNumber="68">Henderson (2002)</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[1063,1484,662,688]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="67" pageNumber="68" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[1063,1484,662,688]" italics="true" pageId="67" pageNumber="68">Snively, Henderson & Phillips (2006)</emphasis>
|
||
</bibRefCitation>
|
||
, and
|
||
<bibRefCitation author="Therrien F & Henderson D & Ruff CB" box="[498,909,702,728]" editor="Carpenter K" journalOrPublisher="Bloomington: Indiana University Press" pageId="67" pageNumber="68" pagination="179 - 237" refId="ref56273" refString="Therrien F, Henderson D, Ruff CB. 2005. Bite me: biomechanical models of theropod mandibles and implications for feeding behavior. In: Carpenter K, ed. The Carnivorous Dinosaurs. Bloomington: Indiana University Press, 179 - 237." title="Bite me: biomechanical models of theropod mandibles and implications for feeding behavior" type="book chapter" volumeTitle="The Carnivorous Dinosaurs" year="2005">
|
||
<emphasis box="[498,909,702,728]" italics="true" pageId="67" pageNumber="68">Therrien, Henderson & Ruff (2005)</emphasis>
|
||
</bibRefCitation>
|
||
mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1324,1393,703,728]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="67" pageNumber="68" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1324,1393,703,728]" italics="true" pageId="67" pageNumber="68">T. rex</emphasis>
|
||
</taxonomicName>
|
||
provides evidence that the ontogenetic transformation of the skull and jaws marks an abrupt change from the juvenile morphotype to a mature morphotype. The skull of juveniles is open framed with a large orbital fenestra and low bone area, which results in low skull strength; also, the bending strength (labiolingually and mesiodistally) of the narrow teeth is low. However, a more powerful bite in juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1063,1131,902,927]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="67" pageNumber="68" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1063,1131,902,927]" italics="true" pageId="67" pageNumber="68">T. rex</emphasis>
|
||
</taxonomicName>
|
||
relative to other theropods of the same size is indicated by the presence of a wide adductor region and fused, vaulted nasals (cf.
|
||
<bibRefCitation author="Snively E & Henderson DM & Phillips DS" box="[619,1026,981,1007]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="67" pageNumber="68" pagination="435 - 454" part="51" refId="ref56086" refString="Snively E, Henderson DM, Phillips DS. 2006. Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics. Acta Palaeontologica Polonica 51: 435 - 454." title="Fused and vaulted nasals of tyrannosaurid dinosaurs: implications for cranial strength and feeding mechanics" type="journal article" year="2006">
|
||
<emphasis box="[619,1026,981,1007]" italics="true" pageId="67" pageNumber="68">Snively, Henderson & Phillips, 2006</emphasis>
|
||
</bibRefCitation>
|
||
). The vertical and relative bending strengths of the mandibular ramus in juveniles are low, which is consistent with the low strength of the cranium.
|
||
</paragraph>
|
||
<paragraph blockId="67.[498,1542,618,1525]" pageId="67" pageNumber="68">The abrupt transition to the mature morphotype (i.e., a tall skull with increase in relative bone area) is seen in subadults, where the nasals have an increased cross section, the nasomaxillary suture acquires a peg-in-socket form, and tooth strength increases (see above for correlates). Although the dorsoventral and relative bending strengths of the mandibular ramus have not been estimated for subadults, they are predicted here to be closer to the high values of adults than to the low values of juveniles.</paragraph>
|
||
<paragraph blockId="67.[498,1542,618,1525]" pageId="67" pageNumber="68">
|
||
In the adult growth categories, the mature morphotype persists largely unchanged aside from continued reduction in orbit ellipticity, inclination, and area; and increase in bone area of the skull, skull strength, maxillary tooth strength, and dorsoventral and relative mandibular strength. Overall, the strength of the craniomandibular skeleton of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1435,1504,1460,1485]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="67" pageNumber="68" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1435,1504,1460,1485]" italics="true" pageId="67" pageNumber="68">T. rex</emphasis>
|
||
</taxonomicName>
|
||
increases throughout adulthood.
|
||
</paragraph>
|
||
<paragraph blockId="67.[498,1542,1570,1921]" box="[498,1146,1570,1600]" pageId="67" pageNumber="68">
|
||
<heading bold="true" box="[498,1146,1570,1600]" fontSize="12" level="2" pageId="67" pageNumber="68" reason="0">
|
||
<emphasis bold="true" box="[498,1146,1570,1600]" pageId="67" pageNumber="68">Ontogenetic change and strain distribution</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="67.[498,1542,1570,1921]" lastBlockId="68.[498,1542,1127,1911]" lastPageId="68" lastPageNumber="69" pageId="67" pageNumber="68">
|
||
Two sets of heat maps of the skull in lateral, dorsal, and ventral views were drafted to show, in relatively high-resolution, the distribution and magnitude of growth changes in each bone (
|
||
<figureCitation box="[636,735,1695,1721]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="67" pageNumber="68">Fig. 26A</figureCitation>
|
||
) and, to reflect a lower-resolution pattern, the frequency in each module (
|
||
<figureCitation box="[604,698,1735,1761]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="67" pageNumber="68">Fig. 26B</figureCitation>
|
||
;
|
||
<tableCitation box="[712,797,1735,1761]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="67" pageNumber="68" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
). In the diagram, the percentage of changes was doubled so that the lightest shades of gray could be seen (i.e., 10% of total changes was rendered as 20% gray, and so on). In the heat map of individual bones, the greatest number of changes are seen from the orbit to the snout, specifically in the lacrimal, and maxilla, jugal, postorbital, frontal, and the dentary (
|
||
<figureCitation box="[938,1037,1894,1921]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="67" pageNumber="68">Fig. 26A</figureCitation>
|
||
). An intermediate number of changes are seen in the nasal, squamosal, quadratojugal, parietal, and braincase, and in the surangular and prearticular of the mandibular ramus (
|
||
<figureCitation box="[1009,1108,1167,1193]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69">Fig. 26A</figureCitation>
|
||
). Finally, the fewest changes of the cranium are seen in the palate and quadrate, and the angular of the lower jaw (
|
||
<figureCitation box="[1427,1525,1206,1232]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69">Fig. 26A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825026" ID-Zenodo-Dep="5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69" startId="68.[524,589,904,926]" targetBox="[498,1541,234,890]" targetPageId="68">
|
||
<paragraph blockId="68.[524,1515,904,1072]" pageId="68" pageNumber="69">
|
||
<emphasis bold="true" pageId="68" pageNumber="69">
|
||
Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="68" pageNumber="69" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" italics="true" pageId="68" pageNumber="69">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="68.[498,1542,1127,1911]" pageId="68" pageNumber="69">
|
||
In the heat map of the modules, the greatest amount of change is seen in the dorsal skull roof, including the nasals, lacrimal, postorbital, and frontals (
|
||
<figureCitation box="[1222,1318,1286,1312]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69">Fig. 26B</figureCitation>
|
||
). An intermediate number of changes is seen in the snout, including the premaxilla, maxilla, jugal, postorbital, and palate (
|
||
<figureCitation box="[772,866,1366,1392]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69">Fig. 26B</figureCitation>
|
||
). Finally, the fewest changes are seen in the suspensorium, parietal, and braincase, and in the mandibular ramus (
|
||
<figureCitation box="[1144,1240,1406,1432]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="68" pageNumber="69">Fig. 26B</figureCitation>
|
||
). In both sets of heat maps, the greatest number of changes occur in the circumorbital region whereas the fewest occur in the suspensorium, parietal, braincase, and postdentary moiety of the mandibular ramus.
|
||
</paragraph>
|
||
<paragraph blockId="68.[498,1542,1127,1911]" pageId="68" pageNumber="69">
|
||
These distributions were compared to two-dimensional models of strain in the cranium of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[530,598,1606,1631]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="68" pageNumber="69" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[530,598,1606,1631]" italics="true" pageId="68" pageNumber="69">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Rayfield EJ" box="[616,779,1605,1631]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="68" pageNumber="69" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[616,779,1605,1631]" italics="true" pageId="68" pageNumber="69">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
) to test the hypothesis that the patterns of growth and strain are no different. Two models of the skull (fused, mobile) were loaded vertically (i.e., biting load) and horizontally (i.e., tearing load); compressive, tensile, and shearing stress were compared (
|
||
<bibRefCitation author="Rayfield EJ" box="[634,797,1725,1751]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="68" pageNumber="69" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[634,797,1725,1751]" italics="true" pageId="68" pageNumber="69">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). In light of recent work showing that the skull of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1399,1468,1725,1750]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="68" pageNumber="69" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1399,1468,1725,1750]" italics="true" pageId="68" pageNumber="69">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was akinetic (
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" box="[608,795,1764,1791]" journalOrPublisher="Anatomical Record" pageId="68" pageNumber="69" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[608,795,1764,1791]" italics="true" pageId="68" pageNumber="69">Cost et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
), the fused skull model results of
|
||
<bibRefCitation author="Rayfield EJ" box="[1197,1372,1764,1791]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="68" pageNumber="69" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[1197,1372,1764,1791]" italics="true" pageId="68" pageNumber="69">Rayfield (2004)</emphasis>
|
||
</bibRefCitation>
|
||
are discussed here.
|
||
</paragraph>
|
||
<paragraph blockId="68.[498,1542,1127,1911]" lastBlockId="69.[498,1542,234,1934]" lastPageId="69" lastPageNumber="70" pageId="68" pageNumber="69">
|
||
During a vertical bite, compressive stress loads the lower half of the skull, especially from the internal antorbital fenestra to the lower temporal bar; tensile stress is highest in the same region (
|
||
<bibRefCitation author="Rayfield EJ" box="[705,868,234,260]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[705,868,234,260]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). In contrast, shear stress is highest above the orbital and internal antorbital fenestra and, to a lesser extent, along the interfenestral strut (
|
||
<bibRefCitation author="Rayfield EJ" box="[508,669,314,340]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[508,669,314,340]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). The ontogenetic heat map for individual bones does not correspond to the compressive or tensile stress distributions, where the region below the orbital- and laterotemporal fenestrae (largely represented by the jugal and quadratojugal) have the lowest proportions of growth changes, with the exception of the high proportion of change seen in the maxilla and lacrimal (
|
||
<figureCitation box="[895,994,473,499]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26A</figureCitation>
|
||
). In contrast, the high proportion of growth changes in the lacrimal and maxilla do reflect the high shear stress seen along the corresponding dorsum of the snout and the interfenestral strut (
|
||
<bibRefCitation author="Rayfield EJ" box="[1251,1413,553,579]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[1251,1413,553,579]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1427,1525,553,579]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="69.[498,1542,234,1934]" pageId="69" pageNumber="70">
|
||
In comparison with the module-based heat map, the region of lowest strain occurs where the greatest amount of change is seen (nasals, lacrimal, postorbital), and the region of highest strain corresponds to the region of an intermediate amount of growth change (maxilla, jugal;
|
||
<figureCitation box="[770,866,712,738]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26B</figureCitation>
|
||
). However, the regions of maximum shear stress do correspond with the region of the greatest growth change, and one of the two regions of lowest shear stress (at the suspensorium), and of tensile stress, does correspond to where the lowest amount of growth changes are seen (
|
||
<figureCitation box="[1064,1160,832,858]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26B</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="69.[498,1542,234,1934]" pageId="69" pageNumber="70">
|
||
During a horizontal tear, the compressive loadings are similar to that of a bite, except that highest loadings are seen adjacent to the loaded maxillary teeth (
|
||
<bibRefCitation author="Rayfield EJ" box="[1319,1482,912,938]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[1319,1482,912,938]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). Tensile stress is similar to that of a bite, concentrated along the lower margin of the snout and orbitotemporal regions (
|
||
<bibRefCitation author="Rayfield EJ" box="[839,1002,991,1018]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[839,1002,991,1018]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). Shear stress is partly similar to the pattern seen in a bite, where stress is highest at the dorsal ramus of the lacrimal, but it is also high below the postorbital bar (
|
||
<bibRefCitation author="Rayfield EJ" box="[812,975,1071,1097]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="69" pageNumber="70" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[812,975,1071,1097]" italics="true" pageId="69" pageNumber="70">Rayfield, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). The ontogenetic heat map does not directly correspond to the stress distributions, aside from the high compressive stress on the maxilla and the shear stress seen on the lacrimal (
|
||
<figureCitation box="[1090,1189,1151,1177]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26A</figureCitation>
|
||
). Therefore, it appears that skull growth did not track stress loads on the skull, whether imposed by biting or tearing. If skull growth was controlled by stress, then higher proportions of change would be seen in the jugal, postorbital, and suspensorium.
|
||
</paragraph>
|
||
<paragraph blockId="69.[498,1542,234,1934]" pageId="69" pageNumber="70">In comparison with the module-based heat map, the pattern of maximum strain and tensile stress, along the subfenestral regions of the maxilla and jugal, do not correspond with the heat map, where the greatest changes are seen along the snout dorsum. In contrast, one of the two regions of maximum shear stress, along the snout dorsum, corresponds to the maximum number of growth changes seen there.</paragraph>
|
||
<paragraph blockId="69.[498,1542,234,1934]" pageId="69" pageNumber="70">
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" box="[530,729,1509,1536]" journalOrPublisher="Anatomical Record" pageId="69" pageNumber="70" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[530,729,1509,1536]" italics="true" pageId="69" pageNumber="70">Cost et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
tested hypotheses of palatal kinesis and strain in the skull of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1457,1526,1510,1535]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="69" pageNumber="70" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1457,1526,1510,1535]" italics="true" pageId="69" pageNumber="70">T. rex</emphasis>
|
||
</taxonomicName>
|
||
using 3-dimensional FEA modeling; the ontogenetic heat maps were compared to their results to test the null hypothesis that the ontogenetic and strain patterns are no different.
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" box="[498,697,1629,1655]" journalOrPublisher="Anatomical Record" pageId="69" pageNumber="70" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[498,697,1629,1655]" italics="true" pageId="69" pageNumber="70">Cost et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
found that strains, in each of the three loading postures (neutral, rostrocaudal, mediolateral) were concentrated along the rostrodorsal margin of the internal antorbital fenestra, palate (including the quadrate), the maxilla adjacent to the palatine, the preorbital bar, base of the postorbital bar, and the lower temporal bar.
|
||
</paragraph>
|
||
<paragraph blockId="69.[498,1542,234,1934]" lastBlockId="70.[498,1542,234,699]" lastPageId="70" lastPageNumber="71" pageId="69" pageNumber="70">
|
||
The ontogenetic heat map for individual bones is consistent with those results in that the highest number of growth changes are seen in the maxilla and lacrimal, but it is inconsistent in that the palate, and the postorbital and lower temporal bars have the lowest proportion of growth changes (
|
||
<figureCitation box="[869,968,1908,1934]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="69" pageNumber="70">Fig. 26A</figureCitation>
|
||
). Therefore, growth changes do not match the strain patterns, indicating that growth does not follow skull loading, aside from the immediate preorbital region, and so a tightly corresponding pattern of remodeling is not seen (
|
||
<figureCitation box="[568,667,314,340]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="70" pageNumber="71">Fig. 26A</figureCitation>
|
||
). These results are consistent with what is seen in the stress models of
|
||
<bibRefCitation author="Rayfield EJ" box="[498,674,354,380]" journalOrPublisher="Proceedings of the Royal Society of London Series B" pageId="70" pageNumber="71" pagination="1451 - 1459" part="271" refId="ref55759" refString="Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London Series B 271 (1547): 1451 - 1459 DOI 10.1098 / rspb. 2004.2755." title="Cranial mechanics and feeding in Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[498,674,354,380]" italics="true" pageId="70" pageNumber="71">Rayfield (2004)</emphasis>
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,234,699]" pageId="70" pageNumber="71">
|
||
In comparison with the module-based heat map, the pattern in lateral view differs, where the regions of maximum strain (subfenestral regions of maxilla and the jugal, and the quadrate) in the model do not correspond to those of the maximum amount of growth changes (snout dorsum). In ventral view, correspondence is seen where the highest strain occurs in the palate and snout, whereas the lowest amount of strain is seen in the braincase (
|
||
<figureCitation box="[626,722,593,619]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="70" pageNumber="71">Fig. 26B</figureCitation>
|
||
). Prima facie, the general noncongruence between growth change distribution and strain distribution indicates that ontogenetic changes were not limited by the requirements for architectural stability and force transmission.
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,744,1932]" box="[498,1417,744,774]" pageId="70" pageNumber="71">
|
||
<heading bold="true" box="[498,1417,744,774]" fontSize="12" level="2" pageId="70" pageNumber="71" reason="0">
|
||
<emphasis bold="true" box="[498,1417,744,774]" pageId="70" pageNumber="71">
|
||
Cephalic musculature lever arm and extension (
|
||
<bibRefCitation author="Molnar RE" box="[1219,1407,744,774]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis bold="true" box="[1219,1407,744,774]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,744,1932]" pageId="70" pageNumber="71">
|
||
<bibRefCitation author="Molnar RE" box="[498,668,789,816]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[498,668,789,816]" italics="true" pageId="70" pageNumber="71">Molnar (2013)</emphasis>
|
||
</bibRefCitation>
|
||
graphed the lever arm and extension of each adductor muscle and the mandibular depressor for juvenile (
|
||
<materialsCitation box="[912,1069,829,856]" collectionCode="CMNH" pageId="70" pageNumber="71" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
) and adult (
|
||
<materialsCitation box="[1214,1451,829,856]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="70" pageNumber="71" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
)
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1467,1535,830,855]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1467,1535,830,855]" italics="true" pageId="70" pageNumber="71">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and adult
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[619,737,870,895]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[619,737,870,895]" italics="true" pageId="70" pageNumber="71">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
(CMN 8506). The anterior pterygoid muscle is excluded from this comparison, as it has been shown by
|
||
<bibRefCitation author="Witmer LM" box="[933,1104,909,935]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="70" pageNumber="71" pagination="1 - 76" part="17" refId="ref56606" refString="Witmer LM. 1997. The evolution of the antorbital cavity of archosaurs: a study in soft-tissue reconstruction in the fossil record with and analysis of the function of pneumaticity. Journal of Vertebrate Paleontology 17 (Suppl. 1): 1 - 76 DOI 10.1080 / 02724634.1997.10011027." title="The evolution of the antorbital cavity of archosaurs: a study in soft-tissue reconstruction in the fossil record with and analysis of the function of pneumaticity" type="journal article" year="1997">
|
||
<emphasis box="[933,1104,909,935]" italics="true" pageId="70" pageNumber="71">Witmer (1997)</emphasis>
|
||
</bibRefCitation>
|
||
that the antorbital fossa was apposed to a paranasal sinus, and not the origin for that muscle.
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,744,1932]" pageId="70" pageNumber="71">
|
||
A marked transition in the lever arm is seen from juvenile to adult in several adductors. When the mouth is closed, the lever arm increases in the posterior mandibular adductor, superficial and medial external mandibular adductor, and the mandibular depressor (
|
||
<bibRefCitation author="Molnar RE" box="[508,665,1108,1134]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[508,665,1108,1134]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). In contrast, that of the deep external mandibular adductor and the pseudotemporal stays the same and that of the dorsal pterygoid decreases (
|
||
<bibRefCitation author="Molnar RE" box="[1370,1526,1148,1174]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1370,1526,1148,1174]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). In
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[530,645,1189,1214]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[530,645,1189,1214]" italics="true" pageId="70" pageNumber="71">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
, the lever arms are approximately intermediate between juvenile and adult for the posterior mandibular adductor, and the superficial and medial external mandibular adductor (
|
||
<bibRefCitation author="Molnar RE" box="[622,779,1268,1294]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[622,779,1268,1294]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,744,1932]" pageId="70" pageNumber="71">
|
||
When the mouth is opened, the lever arm increases from juvenile to adult in the posterior mandibular adductor, superficial and medial external mandibular adductor, and the dorsal pterygoid, whereas that of the deep external mandibular adductor, pseudotemporal, and mandibular depressor stays the same (
|
||
<bibRefCitation author="Molnar RE" box="[1204,1360,1427,1453]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1204,1360,1427,1453]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). In
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1418,1535,1428,1453]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[1418,1535,1428,1453]" italics="true" pageId="70" pageNumber="71">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
, the lever arms are intermediate between juvenile and adult for the superficial and medial external mandibular adductor (
|
||
<bibRefCitation author="Molnar RE" box="[956,1113,1507,1533]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[956,1113,1507,1533]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). If the ontogenetic progression is congruent with the phylogenetic progression, it is predicted that the lever arms in subadult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,567,1587,1612]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,1587,1612]" italics="true" pageId="70" pageNumber="71">T. rex</emphasis>
|
||
</taxonomicName>
|
||
will match what is seen in
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[893,1010,1587,1612]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[893,1010,1587,1612]" italics="true" pageId="70" pageNumber="71">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
for the posterior mandibular adductor, and the superficial and medial external mandibular adductor when the mouth is closed, and the superficial and medial external mandibular adductor when the mouth is agape. The relative magnitude of the lever arm for the muscles is the same in all three exemplars (
|
||
<bibRefCitation author="Molnar RE" box="[508,665,1746,1772]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[508,665,1746,1772]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="70.[498,1542,744,1932]" lastBlockId="71.[498,1542,1236,1581]" lastPageId="71" lastPageNumber="72" pageId="70" pageNumber="71">
|
||
A transition in percentage extension is seen from juvenile to adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1320,1389,1787,1812]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="70" pageNumber="71" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1320,1389,1787,1812]" italics="true" pageId="70" pageNumber="71">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, where an increase occurs in the posterior mandibular adductor, superficial, medial external mandibular adductor, and mandibular depressor (
|
||
<bibRefCitation author="Molnar RE" box="[1081,1236,1865,1892]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1081,1236,1865,1892]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). In contrast, a decrease is seen in the pseudotemporal and the deep external mandibular adductor (
|
||
<bibRefCitation author="Molnar RE" box="[1363,1521,1905,1931]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="70" pageNumber="71" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1363,1521,1905,1931]" italics="true" pageId="70" pageNumber="71">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). The percentage extension in
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[839,957,1237,1262]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[839,957,1237,1262]" italics="true" pageId="71" pageNumber="72">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
is intermediate for the posterior mandibular adductor and the superficial and medial external mandibular adductor (
|
||
<bibRefCitation author="Molnar RE" box="[1347,1504,1275,1302]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="71" pageNumber="72" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1347,1504,1275,1302]" italics="true" pageId="71" pageNumber="72">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
); assuming that the ontogenetic progression will map onto the phylogenetic transition, the expected percentage in subadult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[883,952,1356,1381]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[883,952,1356,1381]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is predicted to be the same as in
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1351,1469,1356,1381]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[1351,1469,1356,1381]" italics="true" pageId="71" pageNumber="72">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
. The percentage extension in
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[839,957,1396,1421]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[839,957,1396,1421]" italics="true" pageId="71" pageNumber="72">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
is lower than in juvenile and adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1380,1448,1396,1421]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1380,1448,1396,1421]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
for the pseudotemporal, deep external mandibular adductor, and the mandibular depressor, indicating that the high extension of those muscles is autapomorphic in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1354,1423,1476,1501]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
(
|
||
<bibRefCitation author="Molnar RE" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="71" pageNumber="72" pagination="177 - 193" refId="ref55179" refString="Molnar RE. 2013. A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 177 - 193." title="A comparative analysis of reconstructed jaw musculature and mechanics of some large theropods" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis italics="true" pageId="71" pageNumber="72">Molnar, 2013</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. It is predicted here that the value in subadults will be intermediate between the values for juvenile and adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[841,910,1555,1580]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[841,910,1555,1580]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, but it will be closer to adults than to juveniles.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825028" ID-Zenodo-Dep="5825028" httpUri="https://zenodo.org/record/5825028/files/figure.png" pageId="71" pageNumber="72" startId="71.[524,589,954,976]" targetBox="[489,1473,226,934]" targetPageId="71">
|
||
<paragraph blockId="71.[524,1516,954,1181]" pageId="71" pageNumber="72">
|
||
<emphasis bold="true" pageId="71" pageNumber="72">
|
||
Figure 27 Comparison of the results of
|
||
<bibRefCitation author="Henderson DM & Snively E." box="[948,1242,954,976]" journalOrPublisher="Proceedings of the Royal Society B" pageId="71" pageNumber="72" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">Henderson & Snively (2004)</bibRefCitation>
|
||
with the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,718,984,1005]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[524,718,984,1005]" italics="true" pageId="71" pageNumber="72">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The rotational inertia (RI) of smaller and progressively distant sister taxa of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[548,602,1013,1034]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[548,602,1013,1034]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
serve as predictive proxies for the RIs of young adult and juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1232,1286,1013,1034]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1232,1286,1013,1034]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Given the larger size of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,577,1042,1063]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[524,577,1042,1063]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
in contrast to non-tyrannosaurine tyrannosaurids, the RI of young adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1266,1319,1042,1063]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1266,1319,1042,1063]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
will almost certainly be more comparable to that of adult
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[875,971,1072,1093]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[875,971,1072,1093]" italics="true" pageId="71" pageNumber="72">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
than to adult
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[1107,1207,1071,1092]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1107,1207,1071,1092]" italics="true" pageId="71" pageNumber="72">A. libratus</emphasis>
|
||
</taxonomicName>
|
||
. The positions of the taxa, aside from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[576,802,1100,1122]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[576,630,1101,1122]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
(
|
||
<materialsCitation box="[642,796,1100,1122]" collectionCode="FMNH" pageId="71" pageNumber="72" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
)
|
||
</taxonomicName>
|
||
, are relative and are not intended to correspond to exact locations along the growth curve. Key to specimens numbered on the growth curve is in Fig. 12.
|
||
</paragraph>
|
||
<paragraph blockId="71.[524,1516,954,1181]" box="[1099,1516,1159,1181]" pageId="71" pageNumber="72">Full-size DOI: 10.7717/peerj.9192/fig-27</paragraph>
|
||
</caption>
|
||
<paragraph blockId="71.[498,1541,1623,1935]" box="[498,1191,1623,1653]" pageId="71" pageNumber="72">
|
||
<heading bold="true" box="[498,1191,1623,1653]" fontSize="12" level="2" pageId="71" pageNumber="72" reason="0">
|
||
<emphasis bold="true" box="[498,1191,1623,1653]" pageId="71" pageNumber="72">
|
||
Rotational inertia (
|
||
<bibRefCitation author="Henderson DM & Snively E." box="[777,1180,1623,1653]" journalOrPublisher="Proceedings of the Royal Society B" pageId="71" pageNumber="72" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">
|
||
<emphasis bold="true" box="[777,1180,1623,1653]" italics="true" pageId="71" pageNumber="72">Henderson & Snively, 2004</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="71.[498,1541,1623,1935]" lastBlockId="72.[498,1507,1236,1303]" lastPageId="72" lastPageNumber="73" pageId="71" pageNumber="72">
|
||
<bibRefCitation author="Henderson DM & Snively E." box="[498,818,1669,1695]" journalOrPublisher="Proceedings of the Royal Society B" pageId="71" pageNumber="72" pagination="S 57 - S 60" part="271" refId="ref54208" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proceedings of the Royal Society B 271: S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[498,818,1669,1695]" italics="true" pageId="71" pageNumber="72">Henderson & Snively (2004)</emphasis>
|
||
</bibRefCitation>
|
||
quantified the rotational inertia (RI) of several tyrannosaurids, including a juvenile and adult of
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[891,1013,1709,1735]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[891,910,1709,1735]" italics="true" pageId="71" pageNumber="72">A</emphasis>
|
||
.
|
||
<emphasis box="[925,1013,1709,1735]" italics="true" pageId="71" pageNumber="72">libratus</emphasis>
|
||
</taxonomicName>
|
||
, an adult of
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1165,1283,1709,1735]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[1165,1185,1710,1735]" italics="true" pageId="71" pageNumber="72">D</emphasis>
|
||
.
|
||
<emphasis box="[1201,1283,1710,1735]" italics="true" pageId="71" pageNumber="72">torosus</emphasis>
|
||
</taxonomicName>
|
||
, and an adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1456,1525,1710,1735]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1456,1525,1710,1735]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Based on mass, it is predicted here that (1) the RI of juvenile
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1227,1296,1749,1774]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1227,1296,1749,1774]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(less than 500 kg) will approximate that of juvenile
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[891,1014,1788,1815]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[891,1014,1788,1815]" italics="true" pageId="71" pageNumber="72">A. libratus</emphasis>
|
||
</taxonomicName>
|
||
, (2)
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1072,1140,1789,1814]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1072,1140,1789,1814]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
in the transition between subadult and adult (~2,800 kg) will be similar to that of adult
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[1217,1338,1828,1855]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1217,1338,1828,1855]" italics="true" pageId="71" pageNumber="72">A. libratus</emphasis>
|
||
</taxonomicName>
|
||
, and (3)
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1444,1511,1829,1854]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1444,1511,1829,1854]" italics="true" pageId="71" pageNumber="72">T. rex</emphasis>
|
||
</taxonomicName>
|
||
in the young adult stage (~3,800 kg; this mass has since been revised to 3,085 kg in
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" journalOrPublisher="PeerJ" pageId="71" pageNumber="72" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="71" pageNumber="72">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
) will be comparable to that of adult
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1062,1180,1909,1934]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="71" pageNumber="72" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[1062,1180,1909,1934]" italics="true" pageId="71" pageNumber="72">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
; ergo, the ontogenetic and phylogenetic trend of tyrannosaurid RI will predict the ontogenetic RI trend of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1438,1507,1237,1262]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1438,1507,1237,1262]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[508,588,1276,1303]" captionStart="Figure 27" captionStartId="71.[524,589,954,976]" captionTargetBox="[489,1473,226,934]" captionTargetId="figure-277@71.[536,1005,247,892]" captionTargetPageId="71" captionText="Figure 27 Comparison of the results of Henderson & Snively (2004) with the growth curve of Tyrannosaurus rex. The rotational inertia (RI) of smaller and progressively distant sister taxa of T. rex serve as predictive proxies for the RIs of young adult and juvenile T. rex. Given the larger size of T. rex in contrast to non-tyrannosaurine tyrannosaurids, the RI of young adult T. rex will almost certainly be more comparable to that of adult D. torosus than to adult A. libratus. The positions of the taxa, aside from T. rex (FMNH PR2081), are relative and are not intended to correspond to exact locations along the growth curve. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-27" figureDoi="http://doi.org/10.5281/zenodo.5825028" httpUri="https://zenodo.org/record/5825028/files/figure.png" pageId="72" pageNumber="73">Fig. 27</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825030" ID-Zenodo-Dep="5825030" httpUri="https://zenodo.org/record/5825030/files/figure.png" pageId="72" pageNumber="73" startId="72.[524,589,955,977]" targetBox="[487,1473,238,936]" targetPageId="72">
|
||
<paragraph blockId="72.[524,1515,954,1181]" pageId="72" pageNumber="73">
|
||
<emphasis bold="true" pageId="72" pageNumber="73">
|
||
Figure 28 Comparison of the results of
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[919,1112,954,976]" journalOrPublisher="PeerJ" pageId="72" pageNumber="73" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis bold="true" box="[919,1112,954,976]" italics="true" pageId="72" pageNumber="73">Snively et al. (2019)</emphasis>
|
||
</bibRefCitation>
|
||
with the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" italics="true" pageId="72" pageNumber="73">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
A comparison of the agility values between
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[977,1031,985,1006]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[977,1031,985,1006]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and other tyrannosaurids. The values for
|
||
<taxonomicName box="[1427,1515,984,1006]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1427,1515,984,1006]" italics="true" pageId="72" pageNumber="73">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[564,658,1014,1035]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[564,658,1014,1035]" italics="true" pageId="72" pageNumber="73">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
serve as predictive proxies for the corresponding values in subadult and young adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1456,1510,1014,1035]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1456,1510,1014,1035]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. The values for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[663,717,1043,1064]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[663,717,1043,1064]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
are calibrated to the growth series, but those of the other taxa are positioned relative to the values seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[728,784,1072,1093]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[728,784,1072,1093]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Inset of the data in table form shows the trends in the data; low values for
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[524,736,1101,1122]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[524,736,1101,1122]" italics="true" pageId="72" pageNumber="73">Albertosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
are in boldface italics. Key to specimens numbered on the growth curve is in Fig. 12.
|
||
<emphasis box="[610,631,1130,1151]" italics="true" pageId="72" pageNumber="73">Al</emphasis>
|
||
,
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[648,862,1130,1151]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[648,862,1130,1151]" italics="true" pageId="72" pageNumber="73">Albertosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<emphasis box="[879,903,1131,1152]" italics="true" pageId="72" pageNumber="73">Dt</emphasis>
|
||
,
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[919,1141,1130,1152]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis box="[919,1141,1130,1152]" italics="true" pageId="72" pageNumber="73">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<emphasis box="[1158,1184,1130,1151]" italics="true" pageId="72" pageNumber="73">Tb</emphasis>
|
||
,
|
||
<taxonomicName box="[1200,1417,1130,1152]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1200,1417,1130,1152]" italics="true" pageId="72" pageNumber="73">Tyrannosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<emphasis box="[1434,1457,1131,1152]" italics="true" pageId="72" pageNumber="73">Tr</emphasis>
|
||
,
|
||
<emphasis italics="true" pageId="72" pageNumber="73">
|
||
Tyrannosaurus
|
||
<taxonomicName baseAuthorityName="FMNH PR" baseAuthorityYear="2081" box="[641,670,1160,1181]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">rex</taxonomicName>
|
||
</emphasis>
|
||
. Full-size DOI: 10.7717/peerj.9192/fig-28
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="72.[498,1542,1347,1738]" box="[498,902,1347,1377]" pageId="72" pageNumber="73">
|
||
<heading bold="true" box="[498,902,1347,1377]" fontSize="12" level="2" pageId="72" pageNumber="73" reason="0">
|
||
<emphasis bold="true" box="[498,902,1347,1377]" pageId="72" pageNumber="73">
|
||
Agility (
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[614,891,1347,1377]" journalOrPublisher="PeerJ" pageId="72" pageNumber="73" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis bold="true" box="[614,891,1347,1377]" italics="true" pageId="72" pageNumber="73">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="72.[498,1542,1347,1738]" pageId="72" pageNumber="73">
|
||
In all tyrannosaurids, including
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[874,942,1394,1419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[874,942,1394,1419]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, agility decreases from the juvenile to adult growth stage (
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[575,792,1433,1459]" journalOrPublisher="PeerJ" pageId="72" pageNumber="73" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[575,792,1433,1459]" italics="true" pageId="72" pageNumber="73">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
), a trend that continues into senescent adulthood (
|
||
<figureCitation box="[1399,1478,1433,1459]" captionStart="Figure 28" captionStartId="72.[524,589,955,977]" captionTargetBox="[487,1473,238,936]" captionTargetId="figure-221@72.[535,1005,247,893]" captionTargetPageId="72" captionText="Figure 28 Comparison of the results of Snively et al. (2019) with the growth curve of Tyrannosaurus rex. A comparison of the agility values between T. rex and other tyrannosaurids. The values for T. bataar and D.torosus serve as predictive proxies for the corresponding values in subadult and young adult T. rex. The values for T. rex are calibrated to the growth series, but those of the other taxa are positioned relative to the values seen in T. rex. Inset of the data in table form shows the trends in the data; low values for Albertosaurus libratus are in boldface italics. Key to specimens numbered on the growth curve is in Fig. 12. Al, Albertosaurus libratus; Dt, Daspletosaurus torosus; Tb, Tyrannosaurus bataar; Tr, Tyr- annosaurus rex. Full-size DOI: 10.7717/peerj.9192/fig-28" figureDoi="http://doi.org/10.5281/zenodo.5825030" httpUri="https://zenodo.org/record/5825030/files/figure.png" pageId="72" pageNumber="73">Fig. 28</figureCitation>
|
||
); however, the adult sample for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[858,927,1473,1498]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[858,927,1473,1498]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
includes only two specimens (
|
||
<materialsCitation box="[1289,1526,1472,1499]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="72" pageNumber="73" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,686,1513,1539]" collectionCode="FMNH" pageId="72" pageNumber="73" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
), so the trend may disappear with additional fossils. Using the mass of other tyrannosaurids as a guide, the transition from high to low agility between the tyrannosaurines is continuous, whereas the values for
|
||
<taxonomicName baseAuthorityName="Snively, Henderson & Phillips" baseAuthorityYear="2006" box="[1123,1244,1592,1619]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis box="[1123,1244,1592,1619]" italics="true" pageId="72" pageNumber="73">A. libratus</emphasis>
|
||
</taxonomicName>
|
||
are slightly lower, but are consistent with the overall trend in reduction (
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[1051,1269,1632,1658]" journalOrPublisher="PeerJ" pageId="72" pageNumber="73" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[1051,1269,1632,1658]" italics="true" pageId="72" pageNumber="73">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1283,1363,1632,1658]" captionStart="Figure 28" captionStartId="72.[524,589,955,977]" captionTargetBox="[487,1473,238,936]" captionTargetId="figure-221@72.[535,1005,247,893]" captionTargetPageId="72" captionText="Figure 28 Comparison of the results of Snively et al. (2019) with the growth curve of Tyrannosaurus rex. A comparison of the agility values between T. rex and other tyrannosaurids. The values for T. bataar and D.torosus serve as predictive proxies for the corresponding values in subadult and young adult T. rex. The values for T. rex are calibrated to the growth series, but those of the other taxa are positioned relative to the values seen in T. rex. Inset of the data in table form shows the trends in the data; low values for Albertosaurus libratus are in boldface italics. Key to specimens numbered on the growth curve is in Fig. 12. Al, Albertosaurus libratus; Dt, Daspletosaurus torosus; Tb, Tyrannosaurus bataar; Tr, Tyr- annosaurus rex. Full-size DOI: 10.7717/peerj.9192/fig-28" figureDoi="http://doi.org/10.5281/zenodo.5825030" httpUri="https://zenodo.org/record/5825030/files/figure.png" pageId="72" pageNumber="73">Fig. 28</figureCitation>
|
||
). Since the extreme growth changes in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[815,882,1673,1698]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[815,882,1673,1698]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
are concentrated in the head, there is no reason to expect an abrupt growth-related decrease of agility in the transition from juvenile to subadult.
|
||
</paragraph>
|
||
<paragraph blockId="72.[498,1542,1782,1935]" box="[498,790,1782,1812]" pageId="72" pageNumber="73">
|
||
<heading bold="true" box="[498,790,1782,1812]" fontSize="12" level="2" pageId="72" pageNumber="73" reason="0">
|
||
<emphasis bold="true" box="[498,790,1782,1812]" pageId="72" pageNumber="73">Suture morphology</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="72.[498,1542,1782,1935]" lastBlockId="73.[498,1541,1221,1447]" lastPageId="73" lastPageNumber="74" pageId="72" pageNumber="73">
|
||
<bibRefCitation author="Herring SW" box="[498,671,1828,1854]" editor="Hanken J & Hall BK" journalOrPublisher="Chicago: University of Chicago Press" pageId="72" pageNumber="73" pagination="153 - 206" refId="ref54243" refString="Herring SW. 1993. Epigenetic and functional influences on skull growth. In: Hanken J, Hall BK, eds. The Skull: Development. Vol. 1. Chicago: University of Chicago Press, 153 - 206." title="Epigenetic and functional influences on skull growth" type="book chapter" volumeTitle="The Skull: Development. Vol. 1" year="1993">
|
||
<emphasis box="[498,671,1828,1854]" italics="true" pageId="72" pageNumber="73">Herring (1993)</emphasis>
|
||
</bibRefCitation>
|
||
stated that, in vertebrates, there
|
||
<emphasis box="[1060,1072,1828,1854]" italics="true" pageId="72" pageNumber="73">“</emphasis>
|
||
seem to be no cases of beveled sutures becoming butt-ended (sensu
|
||
<bibRefCitation author="Moss ML" box="[835,962,1868,1894]" journalOrPublisher="Anatomical Record" pageId="72" pageNumber="73" pagination="569 - 589" part="127" refId="ref55330" refString="Moss ML. 1957. Experimental alteration of sutural area morphology. Anatomical Record 127 (3): 569 - 589 DOI 10.1002 / ar. 1091270307." title="Experimental alteration of sutural area morphology" type="journal article" year="1957">
|
||
<emphasis box="[835,962,1868,1894]" italics="true" pageId="72" pageNumber="73">Moss, 1957</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
<emphasis box="[971,983,1868,1894]" italics="true" pageId="72" pageNumber="73">”</emphasis>
|
||
. However, in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1144,1211,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="72" pageNumber="73" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1144,1211,1869,1894]" italics="true" pageId="72" pageNumber="73">T. rex</emphasis>
|
||
</taxonomicName>
|
||
this extreme transformation is seen at the frontonasal and surangulodentary contacts. The joint surface for the nasal on the frontals in juveniles (e.g.,
|
||
<materialsCitation box="[887,1029,1222,1248]" collectionCode="DDM" pageId="73" pageNumber="74" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
) is a low, inclined facet, whereas in adults (e.g.,
|
||
<materialsCitation box="[641,773,1261,1287]" collectionCode="MOR" pageId="73" pageNumber="74" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
) the joint surface for the nasal is deeply incised and slot-like. The surangulodentary joint surface is a narrow, low angled and shingle-like overlapping contact in juveniles (e.g., BMRP 2002.4.1), whereas in adults (e.g.,
|
||
<materialsCitation box="[1254,1372,1341,1367]" collectionCode="MOR" pageId="73" pageNumber="74" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
), the contact is a wide, coarse, and flattened contact at which both bones are abruptly mediolaterally expanded.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825032" ID-Zenodo-Dep="5825032" httpUri="https://zenodo.org/record/5825032/files/figure.png" pageId="73" pageNumber="74" startId="73.[524,589,954,976]" targetBox="[481,1491,226,936]" targetPageId="73">
|
||
<paragraph blockId="73.[524,1516,953,1151]" pageId="73" pageNumber="74">
|
||
<emphasis bold="true" pageId="73" pageNumber="74">
|
||
Figure 29 Reptile Encephalization Quotients (REQs) of
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" box="[1081,1430,953,975]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="73" pageNumber="74" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis bold="true" box="[1081,1430,953,975]" italics="true" pageId="73" pageNumber="74">Hurlburt, Ridgley & Witmer (2013)</emphasis>
|
||
</bibRefCitation>
|
||
mapped onto the growth curve of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[774,960,983,1004]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="73" pageNumber="74" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[774,960,983,1004]" italics="true" pageId="73" pageNumber="74">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The REQ is based on a brain mass to endocranial volume ratio of 37% and the parenthetical values following the REQs corresponds to the two different body mass estimates, in metric tonnes, from which the REQs were derived (see
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" box="[1162,1483,1041,1063]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="73" pageNumber="74" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1162,1483,1041,1063]" italics="true" pageId="73" pageNumber="74">Hurlburt, Ridgley & Witmer, 2013</emphasis>
|
||
</bibRefCitation>
|
||
for details). Overall, the REQ of the juvenile greatly exceeds that of adults, and the adults show an increasing ontogenetic progression of REQ values, as first reported by
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" box="[1100,1440,1100,1122]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="73" pageNumber="74" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[1100,1440,1100,1122]" italics="true" pageId="73" pageNumber="74">Hurlburt, Ridgley & Witmer (2013)</emphasis>
|
||
</bibRefCitation>
|
||
. Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-29
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="73.[498,1542,1492,1883]" box="[498,971,1492,1522]" pageId="73" pageNumber="74">
|
||
<heading bold="true" box="[498,971,1492,1522]" fontSize="12" level="2" pageId="73" pageNumber="74" reason="0">
|
||
<emphasis bold="true" box="[498,971,1492,1522]" pageId="73" pageNumber="74">Suture closure in the braincase</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="73.[498,1542,1492,1883]" pageId="73" pageNumber="74">
|
||
Although no braincase suture is unambiguously optimized as a synontomorphy, they can be visualized using the Trace function in MacClade (
|
||
<bibRefCitation author="Maddison WP & Maddison DR" box="[1123,1465,1577,1604]" pageId="73" pageNumber="74" publicationUrl="http://macclade.org" refId="ref55064" refString="Maddison WP, Maddison DR. 2005. MacClade 4: analysis of phylogeny and character evolution. Version 4.08 a. Available at http: // macclade. org." title="MacClade 4: analysis of phylogeny and character evolution. Version 4.08 a" type="url" year="2005">
|
||
<emphasis box="[1123,1465,1577,1604]" italics="true" pageId="73" pageNumber="74">Maddison & Maddison, 2005</emphasis>
|
||
</bibRefCitation>
|
||
). The otoccipitobasioccipital suture of the occipital condyle closes first, no later than the start of the young adult category. This is followed by the laterosphenoidoparietal suture and the otoccipitobasioccipital suture of the subcondylar region, which close by the end of the young adult category. The laterosphenoidoprootic suture is closed by the onset of the young adult category, but there is no data for less mature specimens. Finally, neither the laterosphenoidoorbitosphenoid suture or the parietootoccipital suture shows a hierarchical pattern; both tend to remain open.
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,233,703]" box="[498,889,233,263]" pageId="74" pageNumber="75">
|
||
<heading bold="true" box="[498,889,233,263]" fontSize="12" level="2" pageId="74" pageNumber="75" reason="0">
|
||
<emphasis bold="true" box="[498,889,233,263]" pageId="74" pageNumber="75">Encephalization quotients</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,233,703]" pageId="74" pageNumber="75">
|
||
The results of
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" box="[669,1083,279,305]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="74" pageNumber="75" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[669,1083,279,305]" italics="true" pageId="74" pageNumber="75">Hurlburt, Ridgley & Witmer (2013)</emphasis>
|
||
</bibRefCitation>
|
||
were mapped onto the growth curve (
|
||
<figureCitation box="[508,588,318,345]" captionStart="Figure 29" captionStartId="73.[524,589,954,976]" captionTargetBox="[481,1491,226,936]" captionTargetId="figure-199@73.[536,1005,247,892]" captionTargetPageId="73" captionText="Figure 29 Reptile Encephalization Quotients (REQs) of Hurlburt, Ridgley & Witmer (2013) mapped onto the growth curve of Tyrannosaurus rex. The REQ is based on a brain mass to endocranial volume ratio of 37% and the parenthetical values following the REQs corresponds to the two different body mass estimates, in metric tonnes, from which the REQs were derived (see Hurlburt, Ridgley & Witmer, 2013 for details). Overall, the REQ of the juvenile greatly exceeds that of adults, and the adults show an increasing ontogenetic progression of REQ values, as first reported by Hurlburt, Ridgley & Witmer (2013). Key to specimens numbered on the growth curve is in Fig. 12. Full-size DOI: 10.7717/peerj.9192/fig-29" figureDoi="http://doi.org/10.5281/zenodo.5825032" httpUri="https://zenodo.org/record/5825032/files/figure.png" pageId="74" pageNumber="75">Fig. 29</figureCitation>
|
||
), which show a high Reptile Encephalization Quotient (REQ; brain mass to endocast volume ratio of 37%) in a juvenile (
|
||
<materialsCitation box="[1029,1187,358,385]" collectionCode="CMNH" pageId="74" pageNumber="75" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
) that exceeds those of adults. This growth pattern is also seen in
|
||
<taxonomicName authority="(Hurlburt, Ridgley & Witmer, 2013)" baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="74" pageNumber="75" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[906,1190,398,424]" italics="true" pageId="74" pageNumber="75">Alligator mississippiensis</emphasis>
|
||
(
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="74" pageNumber="75" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis italics="true" pageId="74" pageNumber="75">Hurlburt, Ridgley & Witmer, 2013</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Although adult REQs are significantly lower than what is seen in the juvenile, nonetheless an increasing progression in REQ is seen among the three specimens as first hypothesized, based on body size, by
|
||
<bibRefCitation author="Hurlburt GR & Ridgley RC & Witmer LM" box="[941,1355,518,544]" editor="Parrish JM & Molnar RE & Currie PJ & Koppelhus EB" journalOrPublisher="Bloomington: University of Indiana Press" pageId="74" pageNumber="75" pagination="135 - 153" refId="ref54615" refString="Hurlburt GR, Ridgley RC, Witmer LM. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In: Parrish JM, Molnar RE, Currie PJ, Koppelhus EB, eds. Tyrannosaurid Paleobiology. Bloomington: University of Indiana Press, 135 - 153." title="Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">
|
||
<emphasis box="[941,1355,518,544]" italics="true" pageId="74" pageNumber="75">Hurlburt, Ridgley & Witmer (2013)</emphasis>
|
||
</bibRefCitation>
|
||
. It is naïvely predicted here that the reduction in REQ, from the juvenile value to the adult range, will coincide with the transition from juvenile to subadult growth stage as a part of the complex of changes that reshapes the skull from a shallow to a deep morphotype along with corresponding endocranial alteration.
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,748,1657]" box="[498,1201,748,778]" pageId="74" pageNumber="75">
|
||
<heading bold="true" box="[498,1201,748,778]" fontSize="12" level="2" pageId="74" pageNumber="75" reason="0">
|
||
<emphasis bold="true" box="[498,1201,748,778]" pageId="74" pageNumber="75">Congruence between ontogeny and phylogeny</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,748,1657]" pageId="74" pageNumber="75">
|
||
Comparison of the frequency of phylogenetic synontomorphies (ontogenetic characters that are homologous with phylogenetic characters that have been optimized as supporting a node) and nonphylogenetic changes (ontogenetic characters that do not have a phylogenetic homolog) shows that in all cases but one, the nonphylogenetic changes exceed those of phylogenetic changes (
|
||
<figureCitation box="[942,1005,953,980]" captionStart="Figure 5" captionStartId="18.[524,589,692,714]" captionTargetBox="[498,1541,234,679]" captionTargetId="figure-212@18.[497,1128,233,666]" captionTargetPageId="18" captionText="Figure 5 Comparison of the frequency distributions of phylogenetic and nonphylogenetic synontomorphies in the ontogeny of Tyrannosaurus rex. Growth stage is along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and number of synontomorphies is along the y-axis. Phylogenetic characters are in solid bars; nonphylogenetic characters are in hollow bars. The frequency distributions of both sets of data follow the same general pattern, aside from the flatter distribution of the phylogenetic synontomorphies relative to the nonphylogenetic synontomorphies and the reversed pattern seen at growth stages 7 and 8. Both types of changes occur throughout the lifespan of T. rex, indicating that ontogeny is not strictly congruent with phylogeny. Full-size DOI: 10.7717/peerj.9192/fig-5" figureDoi="http://doi.org/10.5281/zenodo.5824982" httpUri="https://zenodo.org/record/5824982/files/figure.png" pageId="74" pageNumber="75">Fig. 5</figureCitation>
|
||
). Overall they share the same peaks, except for a peak at growth stage 4 in the phylogenetic changes (
|
||
<figureCitation box="[1140,1205,993,1020]" captionStart="Figure 5" captionStartId="18.[524,589,692,714]" captionTargetBox="[498,1541,234,679]" captionTargetId="figure-212@18.[497,1128,233,666]" captionTargetPageId="18" captionText="Figure 5 Comparison of the frequency distributions of phylogenetic and nonphylogenetic synontomorphies in the ontogeny of Tyrannosaurus rex. Growth stage is along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and number of synontomorphies is along the y-axis. Phylogenetic characters are in solid bars; nonphylogenetic characters are in hollow bars. The frequency distributions of both sets of data follow the same general pattern, aside from the flatter distribution of the phylogenetic synontomorphies relative to the nonphylogenetic synontomorphies and the reversed pattern seen at growth stages 7 and 8. Both types of changes occur throughout the lifespan of T. rex, indicating that ontogeny is not strictly congruent with phylogeny. Full-size DOI: 10.7717/peerj.9192/fig-5" figureDoi="http://doi.org/10.5281/zenodo.5824982" httpUri="https://zenodo.org/record/5824982/files/figure.png" pageId="74" pageNumber="75">Fig. 5</figureCitation>
|
||
). The corresponding 214-character region of the data matrix shows that relatively immature specimens are coded with the plesiomorphic character states, whereas relatively mature specimens are coded with apomorphic character states. It is therefore reasonable to expect that the ontogenetic character transformations would be congruent,
|
||
<emphasis box="[1205,1308,1153,1178]" italics="true" pageId="74" pageNumber="75">en masse</emphasis>
|
||
, with their phylogenetic counterparts.
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,748,1657]" pageId="74" pageNumber="75">
|
||
However, only 10 of the 329 unambiguously optimized synontomorphies are homologous with the phylogenetic synapomorphies, and a limited congruence with the phylogenetic pattern recovered in
|
||
<bibRefCitation author="Carr TD & Varricchio DJ & Sedlmayr JC & Roberts EM & Moore JR" box="[947,1150,1312,1338]" journalOrPublisher="Scientific Reports" pageId="74" pageNumber="75" pagination="44942" part="7" refId="ref52780" refString="Carr TD, Varricchio DJ, Sedlmayr JC, Roberts EM, Moore JR. 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7 (1): 44942 DOI 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">
|
||
<emphasis box="[947,1150,1312,1338]" italics="true" pageId="74" pageNumber="75">Carr et al. (2017)</emphasis>
|
||
</bibRefCitation>
|
||
is seen (
|
||
<figureCitation box="[1255,1335,1312,1338]" captionStart="Figure 30" captionStartId="75.[524,589,995,1017]" captionTargetBox="[489,1487,224,966]" captionTargetId="graphics-10@75.[534,1446,260,966]" captionTargetPageId="75" captionText="Figure 30 Comparison of recapitulatory synontomorphies of Tyrannosaurus rex with tyrannosauroid phylogeny. Ten unambiguously optimized synontomorphies are congruent with unambiguously optimized synapomorphies of tyrannosauroid phylogeny, providing limited evidence of recapitulation (see text for details). Numbers to the right correspond to the growth stages in Fig. 2. If recapitulation was present, then the growth stage numbers should increase with progressively exclusive clades; that pattern is not seen here. Full-size DOI: 10.7717/peerj.9192/fig-30" figureDoi="http://doi.org/10.5281/zenodo.5825034" httpUri="https://zenodo.org/record/5825034/files/figure.png" pageId="74" pageNumber="75">Fig. 30</figureCitation>
|
||
;
|
||
<tableCitation box="[1350,1451,1312,1338]" captionStart="Table 19" captionStartId="75.[117,172,1271,1292]" captionTargetBox="[108,1525,1430,1831]" captionText="Table 19 Data used in the correlation test between ontogeny and phylogeny for Tyrannosaurus rex. Phylogenetic topology of Carr et al. (2017) was followed for clade ranks. Boldfaced column gives the means for tied ranks (i.e., midranks). The category “derived tyrannosaurines” (i.e., Lythronax, Nanuqsaurus, Teratophoneus) occurs three times given the presence of three ontogenetic characters recovered in the ontogram that are homologous with corresponding synapomorphies. The 12th growth stage occurs twice given that two synontomorphies occr at that growth stage but their homologs occur at different internodes on the phylogenetic hierarchy. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF8FFFE3FF93FB7F3283FA3D" pageId="74" pageNumber="75" tableUuid="DF5C84E2FF8FFFE3FF93FB7F3283FA3D">Table 19</tableCitation>
|
||
). A Shapiro
|
||
<emphasis box="[616,631,1353,1378]" italics="true" pageId="74" pageNumber="75">–</emphasis>
|
||
Wilk test found that the ranked growth data and clade rank data are normally distributed (
|
||
<emphasis box="[642,656,1393,1418]" italics="true" pageId="74" pageNumber="75">p</emphasis>
|
||
= 0.690 and 0.767, respectively). A Spearman-rank correlation test results in a nonsignificant (
|
||
<emphasis box="[705,719,1432,1457]" italics="true" pageId="74" pageNumber="75">p</emphasis>
|
||
= 0.693) correlation coefficient (
|
||
<emphasis box="[1108,1119,1432,1457]" italics="true" pageId="74" pageNumber="75">
|
||
<emphasis box="[1119,1129,1444,1460]" italics="true" pageId="74" pageNumber="75">
|
||
<subScript attach="left" box="[1119,1129,1444,1460]" fontSize="7" pageId="74" pageNumber="75">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= −0.143;
|
||
<tableCitation box="[1258,1359,1431,1458]" captionStart="Table 19" captionStartId="75.[117,172,1271,1292]" captionTargetBox="[108,1525,1430,1831]" captionText="Table 19 Data used in the correlation test between ontogeny and phylogeny for Tyrannosaurus rex. Phylogenetic topology of Carr et al. (2017) was followed for clade ranks. Boldfaced column gives the means for tied ranks (i.e., midranks). The category “derived tyrannosaurines” (i.e., Lythronax, Nanuqsaurus, Teratophoneus) occurs three times given the presence of three ontogenetic characters recovered in the ontogram that are homologous with corresponding synapomorphies. The 12th growth stage occurs twice given that two synontomorphies occr at that growth stage but their homologs occur at different internodes on the phylogenetic hierarchy. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF8FFFE3FF93FB7F3283FA3D" pageId="74" pageNumber="75" tableUuid="DF5C84E2FF8FFFE3FF93FB7F3283FA3D">Table 19</tableCitation>
|
||
;
|
||
<figureCitation box="[1374,1454,1431,1458]" captionStart="Figure 31" captionStartId="76.[524,589,656,678]" captionTargetBox="[502,1296,237,624]" captionTargetId="figure-415@76.[497,1066,233,629]" captionTargetPageId="76" captionText="Figure 31 Bivariate scatterplot showing the test of ontogenetic recapitulation of phylogenetic novelties in Tyrannosaurus rex. Growth stage rank (increases away from the origin) is along the x-axis; clade rank (increases away from the origin) is along the y-axis. If recapitulation is present, then the ranks will increase montonically from the origin. A recapitulatory pattern is not seen in T. rex; see text for details. Full-size DOI: 10.7717/peerj.9192/fig-31" figureDoi="http://doi.org/10.5281/zenodo.5825036" httpUri="https://zenodo.org/record/5825036/files/figure.png" pageId="74" pageNumber="75">Fig. 31</figureCitation>
|
||
) between growth stage rank and clade rank. The phylogenetic context of the conflicting data are illustrated in
|
||
<figureCitation box="[698,777,1511,1537]" captionStart="Figure 30" captionStartId="75.[524,589,995,1017]" captionTargetBox="[489,1487,224,966]" captionTargetId="graphics-10@75.[534,1446,260,966]" captionTargetPageId="75" captionText="Figure 30 Comparison of recapitulatory synontomorphies of Tyrannosaurus rex with tyrannosauroid phylogeny. Ten unambiguously optimized synontomorphies are congruent with unambiguously optimized synapomorphies of tyrannosauroid phylogeny, providing limited evidence of recapitulation (see text for details). Numbers to the right correspond to the growth stages in Fig. 2. If recapitulation was present, then the growth stage numbers should increase with progressively exclusive clades; that pattern is not seen here. Full-size DOI: 10.7717/peerj.9192/fig-30" figureDoi="http://doi.org/10.5281/zenodo.5825034" httpUri="https://zenodo.org/record/5825034/files/figure.png" pageId="74" pageNumber="75">Fig. 30</figureCitation>
|
||
. It is possible that this result is an artifact of the relatively poorly sampled juvenile and subadult growth categories, which indicates that the discovery of new specimens in those categories of maturity will simultaneously increase the frequency and congruence of phylogenetically homologous character changes.
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,1702,1934]" box="[498,786,1702,1732]" pageId="74" pageNumber="75">
|
||
<heading bold="true" box="[498,786,1702,1732]" fontSize="12" level="2" pageId="74" pageNumber="75" reason="0">
|
||
<emphasis bold="true" box="[498,786,1702,1732]" pageId="74" pageNumber="75">Individual variation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="74.[498,1542,1702,1934]" lastBlockId="76.[498,1542,850,1914]" lastPageId="76" lastPageNumber="77" pageId="74" pageNumber="75">
|
||
By growth category, the frequency of individual variation in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1199,1266,1748,1773]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="74" pageNumber="75" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1199,1266,1748,1773]" italics="true" pageId="74" pageNumber="75">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is, in descending order: adult (151), young adult (98), juvenile (22), senescent adult (19), subadult (6). The low amount of variation at the subadult category almost certainly results from the fact that the growth stage is represented by three incomplete specimens (
|
||
<materialsCitation box="[1252,1440,1867,1893]" collectionCode="BMRP" pageId="74" pageNumber="75" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="LACM" pageId="74" pageNumber="75" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
,
|
||
<materialsCitation box="[581,726,1907,1933]" collectionCode="RSM" pageId="74" pageNumber="75" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
). Nearly twice as much variation is seen in adults than is seen in young adults; given the completeness of specimens in these growth stages, this difference almost certainly approximates the real pattern. In nearly every skull module, the number of changes increases with ascending ontogenetic categories (
|
||
<tableCitation box="[1198,1284,930,956]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="76" pageNumber="77" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
), which is evidence of extensive remodeling throughout the phases of life that correspond to sexual maturity.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825034" ID-Zenodo-Dep="5825034" httpUri="https://zenodo.org/record/5825034/files/figure.png" pageId="75" pageNumber="76" startId="75.[524,589,995,1017]" targetBox="[489,1487,224,966]" targetPageId="75">
|
||
<paragraph pageId="75" pageNumber="76">
|
||
<emphasis bold="true" pageId="75" pageNumber="76">
|
||
Figure 30 Comparison of recapitulatory synontomorphies of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1127,1310,995,1016]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1127,1310,995,1016]" italics="true" pageId="75" pageNumber="76">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
with tyrannosauroid phylogeny.
|
||
</emphasis>
|
||
Ten unambiguously optimized synontomorphies are congruent with unambiguously optimized synapomorphies of tyrannosauroid phylogeny, providing limited evidence of recapitulation (see text for details). Numbers to the right correspond to the growth stages in Fig. 2. If recapitulation was present, then the growth stage numbers should increase with progressively exclusive clades; that pattern is not seen here. Full-size DOI: 10.7717/peerj.9192/fig-30
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FF8FFFE3FF93FB7F3283FA3D" httpUri="http://table.plazi.org/id/DF5C84E2FF8FFFE3FF93FB7F3283FA3D" pageId="75" pageNumber="76" startId="75.[117,172,1271,1292]" targetBox="[108,1525,1430,1831]" targetIsTable="true" targetPageId="75">
|
||
<paragraph blockId="75.[117,1515,1270,1409]" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[117,1104,1270,1292]" pageId="75" pageNumber="76">
|
||
Table 19 Data used in the correlation test between ontogeny and phylogeny for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[913,1099,1271,1292]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[913,1099,1271,1292]" italics="true" pageId="75" pageNumber="76">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Phylogenetic topology of
|
||
<bibRefCitation author="Carr TD & Varricchio DJ & Sedlmayr JC & Roberts EM & Moore JR" box="[1353,1515,1270,1292]" journalOrPublisher="Scientific Reports" pageId="75" pageNumber="76" pagination="44942" part="7" refId="ref52780" refString="Carr TD, Varricchio DJ, Sedlmayr JC, Roberts EM, Moore JR. 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7 (1): 44942 DOI 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">
|
||
<emphasis box="[1353,1515,1270,1292]" italics="true" pageId="75" pageNumber="76">Carr et al. (2017)</emphasis>
|
||
</bibRefCitation>
|
||
was followed for clade ranks. Boldfaced column gives the means for tied ranks (i.e., midranks). The category
|
||
<emphasis box="[1260,1269,1300,1322]" italics="true" pageId="75" pageNumber="76">“</emphasis>
|
||
derived tyrannosaurines
|
||
<emphasis box="[1506,1515,1300,1322]" italics="true" pageId="75" pageNumber="76">”</emphasis>
|
||
(i.e.,
|
||
<emphasis box="[163,261,1329,1350]" italics="true" pageId="75" pageNumber="76">Lythronax</emphasis>
|
||
,
|
||
<emphasis box="[272,399,1330,1351]" italics="true" pageId="75" pageNumber="76">Nanuqsaurus</emphasis>
|
||
,
|
||
<emphasis box="[409,549,1329,1350]" italics="true" pageId="75" pageNumber="76">Teratophoneus</emphasis>
|
||
) occurs three times given the presence of three ontogenetic characters recovered in the ontogram that are homologous with corresponding synapomorphies. The 12th growth stage occurs twice given that two synontomorphies occr at that growth stage but their homologs occur at different internodes on the phylogenetic hierarchy. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="75" pageNumber="76">
|
||
<table box="[108,1525,1430,1831]" gridcols="6" gridrows="11" pageId="75" pageNumber="76">
|
||
<tr box="[108,1525,1430,1452]" gridrow="0" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1430,1452]" gridcol="0" gridrow="0" pageId="75" pageNumber="76">Clade</th>
|
||
<th box="[586,696,1430,1452]" gridcol="1" gridrow="0" pageId="75" pageNumber="76">Clade rank</th>
|
||
<th box="[753,912,1430,1452]" gridcol="2" gridrow="0" pageId="75" pageNumber="76">Clade midranks</th>
|
||
<th box="[969,1102,1430,1452]" gridcol="3" gridrow="0" pageId="75" pageNumber="76">Growth stage</th>
|
||
<th box="[1159,1289,1430,1452]" gridcol="4" gridrow="0" pageId="75" pageNumber="76">Growth rank</th>
|
||
<th box="[1346,1525,1430,1452]" gridcol="5" gridrow="0" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1525,1430,1452]" pageId="75" pageNumber="76">Growth midranks</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1525,1471,1493]" gridrow="1" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1471,1493]" gridcol="0" gridrow="1" pageId="75" pageNumber="76">
|
||
<emphasis box="[108,117,1471,1493]" italics="true" pageId="75" pageNumber="76">“</emphasis>
|
||
Stokesosaurids
|
||
<emphasis box="[257,266,1471,1493]" italics="true" pageId="75" pageNumber="76">”</emphasis>
|
||
+ derived tyrannosauroids
|
||
</th>
|
||
<td box="[586,696,1471,1493]" gridcol="1" gridrow="1" pageId="75" pageNumber="76">1</td>
|
||
<td box="[753,912,1471,1493]" gridcol="2" gridrow="1" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1471,1492]" pageId="75" pageNumber="76">1</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1471,1493]" gridcol="3" gridrow="1" pageId="75" pageNumber="76">9</td>
|
||
<td box="[1159,1289,1471,1493]" gridcol="4" gridrow="1" pageId="75" pageNumber="76">7</td>
|
||
<td box="[1346,1525,1471,1493]" gridcol="5" gridrow="1" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1471,1492]" pageId="75" pageNumber="76">7</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1509,1531]" gridrow="2" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1509,1531]" gridcol="0" gridrow="2" pageId="75" pageNumber="76">
|
||
<emphasis box="[108,250,1509,1530]" italics="true" pageId="75" pageNumber="76">Xiongguanlong</emphasis>
|
||
+ derived tyrannosauroids
|
||
</th>
|
||
<td box="[586,696,1509,1531]" gridcol="1" gridrow="2" pageId="75" pageNumber="76">2</td>
|
||
<td box="[753,912,1509,1531]" gridcol="2" gridrow="2" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1509,1530]" pageId="75" pageNumber="76">2</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1509,1531]" gridcol="3" gridrow="2" pageId="75" pageNumber="76">18</td>
|
||
<td box="[1159,1289,1509,1531]" gridcol="4" gridrow="2" pageId="75" pageNumber="76">10</td>
|
||
<td box="[1346,1525,1509,1531]" gridcol="5" gridrow="2" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1370,1509,1530]" pageId="75" pageNumber="76">10</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1546,1568]" gridrow="3" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1546,1568]" gridcol="0" gridrow="3" pageId="75" pageNumber="76">
|
||
<emphasis box="[108,240,1546,1567]" italics="true" pageId="75" pageNumber="76">Bistahieversor</emphasis>
|
||
+ derived tyrannosauroids
|
||
</th>
|
||
<td box="[586,696,1546,1568]" gridcol="1" gridrow="3" pageId="75" pageNumber="76">3</td>
|
||
<td box="[753,912,1546,1568]" gridcol="2" gridrow="3" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1546,1567]" pageId="75" pageNumber="76">3</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1546,1568]" gridcol="3" gridrow="3" pageId="75" pageNumber="76">5</td>
|
||
<td box="[1159,1289,1546,1568]" gridcol="4" gridrow="3" pageId="75" pageNumber="76">2</td>
|
||
<td box="[1346,1525,1546,1568]" gridcol="5" gridrow="3" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1546,1567]" pageId="75" pageNumber="76">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1584,1606]" gridrow="4" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1584,1606]" gridcol="0" gridrow="4" pageId="75" pageNumber="76">
|
||
<taxonomicName box="[108,270,1584,1606]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
</th>
|
||
<td box="[586,696,1584,1606]" gridcol="1" gridrow="4" pageId="75" pageNumber="76">4</td>
|
||
<td box="[753,912,1584,1606]" gridcol="2" gridrow="4" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1584,1605]" pageId="75" pageNumber="76">4</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1584,1606]" gridcol="3" gridrow="4" pageId="75" pageNumber="76">8</td>
|
||
<td box="[1159,1289,1584,1606]" gridcol="4" gridrow="4" pageId="75" pageNumber="76">4</td>
|
||
<td box="[1346,1525,1584,1606]" gridcol="5" gridrow="4" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1584,1605]" pageId="75" pageNumber="76">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1621,1644]" gridrow="5" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1621,1644]" gridcol="0" gridrow="5" pageId="75" pageNumber="76">derived tyrannosaurines</th>
|
||
<td box="[586,696,1621,1644]" gridcol="1" gridrow="5" pageId="75" pageNumber="76">5</td>
|
||
<td box="[753,912,1621,1644]" gridcol="2" gridrow="5" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1621,1642]" pageId="75" pageNumber="76">6</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1621,1644]" gridcol="3" gridrow="5" pageId="75" pageNumber="76">3</td>
|
||
<td box="[1159,1289,1621,1644]" gridcol="4" gridrow="5" pageId="75" pageNumber="76">1</td>
|
||
<td box="[1346,1525,1621,1644]" gridcol="5" gridrow="5" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1621,1642]" pageId="75" pageNumber="76">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1659,1681]" gridrow="6" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1659,1681]" gridcol="0" gridrow="6" pageId="75" pageNumber="76">derived tyrannosaurines</th>
|
||
<td box="[586,696,1659,1681]" gridcol="1" gridrow="6" pageId="75" pageNumber="76">6</td>
|
||
<td box="[753,912,1659,1681]" gridcol="2" gridrow="6" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1659,1680]" pageId="75" pageNumber="76">6</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1659,1681]" gridcol="3" gridrow="6" pageId="75" pageNumber="76">7</td>
|
||
<td box="[1159,1289,1659,1681]" gridcol="4" gridrow="6" pageId="75" pageNumber="76">3</td>
|
||
<td box="[1346,1525,1659,1681]" gridcol="5" gridrow="6" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1659,1680]" pageId="75" pageNumber="76">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1697,1719]" gridrow="7" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1697,1719]" gridcol="0" gridrow="7" pageId="75" pageNumber="76">derived tyrannosaurines</th>
|
||
<td box="[586,696,1697,1719]" gridcol="1" gridrow="7" pageId="75" pageNumber="76">7</td>
|
||
<td box="[753,912,1697,1719]" gridcol="2" gridrow="7" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1697,1718]" pageId="75" pageNumber="76">6</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1697,1719]" gridcol="3" gridrow="7" pageId="75" pageNumber="76">12</td>
|
||
<td box="[1159,1289,1697,1719]" gridcol="4" gridrow="7" pageId="75" pageNumber="76">8</td>
|
||
<td box="[1346,1525,1697,1719]" gridcol="5" gridrow="7" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1376,1697,1718]" pageId="75" pageNumber="76">8.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1734,1756]" gridrow="8" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1734,1756]" gridcol="0" gridrow="8" pageId="75" pageNumber="76">
|
||
<taxonomicName authorityName="Russell" authorityYear="1970" box="[108,251,1734,1755]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="genus">
|
||
<emphasis box="[108,251,1734,1755]" italics="true" pageId="75" pageNumber="76">Daspletosaurus</emphasis>
|
||
</taxonomicName>
|
||
+ advanced tyrannosaurines
|
||
</th>
|
||
<td box="[586,696,1734,1756]" gridcol="1" gridrow="8" pageId="75" pageNumber="76">8</td>
|
||
<td box="[753,912,1734,1756]" gridcol="2" gridrow="8" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1734,1755]" pageId="75" pageNumber="76">8</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1734,1756]" gridcol="3" gridrow="8" pageId="75" pageNumber="76">12</td>
|
||
<td box="[1159,1289,1734,1756]" gridcol="4" gridrow="8" pageId="75" pageNumber="76">9</td>
|
||
<td box="[1346,1525,1734,1756]" gridcol="5" gridrow="8" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1376,1734,1755]" pageId="75" pageNumber="76">8.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1772,1794]" gridrow="9" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1772,1794]" gridcol="0" gridrow="9" pageId="75" pageNumber="76">
|
||
<emphasis box="[108,284,1772,1793]" italics="true" pageId="75" pageNumber="76">Zhuchengtyrannus</emphasis>
|
||
+
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[312,455,1773,1794]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="genus">
|
||
<emphasis box="[312,455,1773,1794]" italics="true" pageId="75" pageNumber="76">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td box="[586,696,1772,1794]" gridcol="1" gridrow="9" pageId="75" pageNumber="76">9</td>
|
||
<td box="[753,912,1772,1794]" gridcol="2" gridrow="9" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,765,1772,1793]" pageId="75" pageNumber="76">9</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1772,1794]" gridcol="3" gridrow="9" pageId="75" pageNumber="76">8</td>
|
||
<td box="[1159,1289,1772,1794]" gridcol="4" gridrow="9" pageId="75" pageNumber="76">5</td>
|
||
<td box="[1346,1525,1772,1794]" gridcol="5" gridrow="9" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1772,1793]" pageId="75" pageNumber="76">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[108,1525,1809,1831]" gridrow="10" pageId="75" pageNumber="76">
|
||
<th box="[108,529,1809,1831]" gridcol="0" gridrow="10" pageId="75" pageNumber="76">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[108,251,1810,1831]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="75" pageNumber="76" phylum="Chordata" rank="genus">
|
||
<emphasis box="[108,251,1810,1831]" italics="true" pageId="75" pageNumber="76">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td box="[586,696,1809,1831]" gridcol="1" gridrow="10" pageId="75" pageNumber="76">10</td>
|
||
<td box="[753,912,1809,1831]" gridcol="2" gridrow="10" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[753,777,1809,1830]" pageId="75" pageNumber="76">10</emphasis>
|
||
</td>
|
||
<td box="[969,1102,1809,1831]" gridcol="3" gridrow="10" pageId="75" pageNumber="76">8</td>
|
||
<td box="[1159,1289,1809,1831]" gridcol="4" gridrow="10" pageId="75" pageNumber="76">6</td>
|
||
<td box="[1346,1525,1809,1831]" gridcol="5" gridrow="10" pageId="75" pageNumber="76">
|
||
<emphasis bold="true" box="[1346,1358,1810,1831]" pageId="75" pageNumber="76">5</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825036" ID-Zenodo-Dep="5825036" httpUri="https://zenodo.org/record/5825036/files/figure.png" pageId="76" pageNumber="77" startId="76.[524,589,656,678]" targetBox="[502,1296,237,624]" targetPageId="76">
|
||
<paragraph blockId="76.[524,1515,656,795]" pageId="76" pageNumber="77">
|
||
<emphasis bold="true" pageId="76" pageNumber="77">
|
||
Figure 31 Bivariate scatterplot showing the test of ontogenetic recapitulation of phylogenetic novelties in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[653,844,686,707]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="76" pageNumber="77" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[653,844,686,707]" italics="true" pageId="76" pageNumber="77">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Growth stage rank (increases away from the origin) is along the
|
||
<emphasis box="[524,535,715,736]" italics="true" pageId="76" pageNumber="77">x</emphasis>
|
||
-axis; clade rank (increases away from the origin) is along the
|
||
<emphasis box="[1114,1124,715,736]" italics="true" pageId="76" pageNumber="77">y</emphasis>
|
||
-axis. If recapitulation is present, then the ranks will increase montonically from the origin. A recapitulatory pattern is not seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1349,1403,744,765]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="76" pageNumber="77" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1349,1403,744,765]" italics="true" pageId="76" pageNumber="77">T. rex</emphasis>
|
||
</taxonomicName>
|
||
; see text for details. Full-size DOI: 10.7717/peerj.9192/fig-31
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="76.[498,1542,850,1914]" pageId="76" pageNumber="77">
|
||
In comparison with the frequency of synontomorphies at each node, with two exceptions, individual variation outnumbers synontomorphies from the young adult category forward (
|
||
<figureCitation box="[712,775,1089,1115]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="76" pageNumber="77">Fig. 2</figureCitation>
|
||
). The correlation between the individual variation on each branch and the number of synontomorphies at each node was assessed under two correlation tests (
|
||
<tableCitation box="[565,664,1169,1195]" captionStart="Table 20" captionStartId="77.[525,580,248,269]" captionTargetBox="[516,1524,378,1628]" captionText="Table 20 Summary of data used in the Spearman correlation test of individual variation per specimen per node and number of synontomorphies per node for Tyrannnosaurus rex. The rows are organized by node, followed by the sequence of number of synontomorphies and individual variation by number, rank, and midranks. Boldfaced columns indicate values used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF89FFE5FDEBFF7032B2FED9" pageId="76" pageNumber="77" tableUuid="DF5C84E2FF89FFE5FDEBFF7032B2FED9">Table 20</tableCitation>
|
||
). The raw data are skewed to right and were run under a Pearson correlation test that obtained a nonsignificant (
|
||
<emphasis box="[919,933,1209,1234]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.559) correlation (
|
||
<emphasis box="[1194,1205,1209,1234]" italics="true" pageId="76" pageNumber="77">r</emphasis>
|
||
= −0.109). The skewness was reduced upon converting the data to ranks (
|
||
<tableCitation box="[1073,1175,1249,1275]" captionStart="Table 20" captionStartId="77.[525,580,248,269]" captionTargetBox="[516,1524,378,1628]" captionText="Table 20 Summary of data used in the Spearman correlation test of individual variation per specimen per node and number of synontomorphies per node for Tyrannnosaurus rex. The rows are organized by node, followed by the sequence of number of synontomorphies and individual variation by number, rank, and midranks. Boldfaced columns indicate values used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF89FFE5FDEBFF7032B2FED9" pageId="76" pageNumber="77" tableUuid="DF5C84E2FF89FFE5FDEBFF7032B2FED9">Table 20</tableCitation>
|
||
), and a Shapiro
|
||
<emphasis box="[1364,1379,1249,1274]" italics="true" pageId="76" pageNumber="77">–</emphasis>
|
||
Wilk test found that the ranked data are not normally distributed for the individual variation ranks (
|
||
<emphasis box="[581,595,1329,1354]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.015), whereas the data for the synontomorphy ranks are normally distributed (
|
||
<emphasis box="[644,658,1369,1394]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.144). A Spearman rank correlation test was run, which returned a nonsignificant (
|
||
<emphasis box="[683,697,1409,1434]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.695) coefficient (
|
||
<emphasis box="[951,962,1409,1434]" italics="true" pageId="76" pageNumber="77">
|
||
<emphasis box="[962,972,1420,1436]" italics="true" pageId="76" pageNumber="77">
|
||
<subScript attach="left" box="[962,972,1420,1436]" fontSize="7" pageId="76" pageNumber="77">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= −0.073) (
|
||
<figureCitation box="[1114,1193,1408,1434]" captionStart="Figure 32" captionStartId="78.[524,589,656,678]" captionTargetBox="[500,1298,234,623]" captionTargetId="figure-176@78.[497,1099,233,629]" captionTargetPageId="78" captionText="Figure 32 Bivariate scatterplot showing the congruence between individual variation per specimen per node compared with the number of unambiguously optimized synontomorphies per node in Tyrannosaurus rex. The number of synontomorphies per node are along the x-axis; the amount of individual variation (i.e., unambiguously optimized character states per branch) is along the y-axis. Both values increase away from the origin. If the amount of individual variation is controlled by the number of synontomorphies per node, then the variables should increase monotonically. In this case, no congruence is seen between the variables in T. rex. Full-size DOI: 10.7717/peerj.9192/fig-32" figureDoi="http://doi.org/10.5281/zenodo.5825038" httpUri="https://zenodo.org/record/5825038/files/figure.png" pageId="76" pageNumber="77">Fig. 32</figureCitation>
|
||
;
|
||
<tableCitation box="[1208,1310,1408,1434]" captionStart="Table 20" captionStartId="77.[525,580,248,269]" captionTargetBox="[516,1524,378,1628]" captionText="Table 20 Summary of data used in the Spearman correlation test of individual variation per specimen per node and number of synontomorphies per node for Tyrannnosaurus rex. The rows are organized by node, followed by the sequence of number of synontomorphies and individual variation by number, rank, and midranks. Boldfaced columns indicate values used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF89FFE5FDEBFF7032B2FED9" pageId="76" pageNumber="77" tableUuid="DF5C84E2FF89FFE5FDEBFF7032B2FED9">Table 20</tableCitation>
|
||
). Therefore, the number of synontomorphies does not influence the amount of individual variation. A lower-resolution Spearman rank correlation test was done, at the level of growth category (
|
||
<tableCitation box="[615,703,1527,1554]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="76" pageNumber="77" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
), which also resulted in a nonsignificant (
|
||
<emphasis box="[1201,1215,1528,1553]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.397) correlation coefficient (
|
||
<emphasis box="[637,648,1568,1593]" italics="true" pageId="76" pageNumber="77">
|
||
<emphasis box="[647,657,1580,1596]" italics="true" pageId="76" pageNumber="77">
|
||
<subScript attach="left" box="[647,657,1580,1596]" fontSize="7" pageId="76" pageNumber="77">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.429), showing there is no correlation at that low level of comparison.
|
||
</paragraph>
|
||
<paragraph blockId="76.[498,1542,850,1914]" lastBlockId="78.[498,1542,1426,1931]" lastPageId="78" lastPageNumber="79" pageId="76" pageNumber="77">
|
||
In terms of craniomandibular modules, the frequency of individual variation, in descending order, is: dorsum of snout (100), sides of snout (57), mandibular ramus (42), braincase (29), suspensorium (23), and parietal (23) (
|
||
<tableCitation box="[1115,1200,1687,1713]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="76" pageNumber="77" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
; Data S5). As such, no region of the skull is exempt from individual variation. A second correlation test was run to compare the amount of synontomorphies with individual variation in each module (
|
||
<tableCitation box="[508,610,1806,1833]" captionStart="Table 21" captionStartId="78.[525,580,927,948]" captionTargetBox="[516,1525,1056,1366]" captionText="Table 21 Summary of the number of synontomorphies and individual variation per craniomandibular module. Raw and corrected data used in the correlation test between the number of synontomorphies in each skull module with the amount of individual variation per module. Boldface indicates the ranks used in the correlation test." httpUri="http://table.plazi.org/id/DF5C84E2FF8AFFE6FDEBFC1734C8FBB0" pageId="76" pageNumber="77" tableUuid="DF5C84E2FF8AFFE6FDEBFC1734C8FBB0">Table 21</tableCitation>
|
||
); Shapiro
|
||
<emphasis box="[725,740,1807,1832]" italics="true" pageId="76" pageNumber="77">–</emphasis>
|
||
Wilk normality tests found that the synontomorphy data and the individual variation data are normally distributed (
|
||
<emphasis box="[1100,1114,1847,1872]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.961, 0.537, respectively). The correlation test resulted in a significant (
|
||
<emphasis box="[1018,1032,1887,1912]" italics="true" pageId="76" pageNumber="77">p</emphasis>
|
||
= 0.008) correlation coefficient (
|
||
<emphasis box="[1412,1423,1887,1912]" italics="true" pageId="76" pageNumber="77">
|
||
<emphasis box="[1422,1432,1898,1914]" italics="true" pageId="76" pageNumber="77">
|
||
<subScript attach="left" box="[1422,1432,1898,1914]" fontSize="7" pageId="76" pageNumber="77">rS</subScript>
|
||
</emphasis>
|
||
</emphasis>
|
||
= 0.928). The similar amount of individual variation and number of synontomorphies in each skull module suggests that the number of unambiguously optimized characters are dependent upon the proportion of characters that is scored for each module in the data matrix. Indeed, a tally from the character list (Data S1) shows the frequency of characters per module, in descending order: snout and palate (399), skull roof (334), mandible (141), suspensorium (140), braincase (85), and parietal (25). The distribution broadly matches that of the individual variation and synontomorphies, except for the reversal in order of the snout and skull roof modules, and the suspensorium and braincase modules.
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FF89FFE5FDEBFF7032B2FED9" httpUri="http://table.plazi.org/id/DF5C84E2FF89FFE5FDEBFF7032B2FED9" pageId="77" pageNumber="78" startId="77.[525,580,248,269]" targetBox="[516,1524,378,1628]" targetIsTable="true" targetPageId="77">
|
||
<paragraph blockId="77.[525,1515,248,358]" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" pageId="77" pageNumber="78">
|
||
Table 20 Summary of data used in the Spearman correlation test of individual variation per specimen per node and number of synontomorphies per node for
|
||
<emphasis bold="true" box="[1208,1409,278,299]" italics="true" pageId="77" pageNumber="78">
|
||
Tyrannnosaurus
|
||
<taxonomicName baseAuthorityName="FMNH PR" baseAuthorityYear="2081" box="[1378,1409,278,299]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="77" pageNumber="78" phylum="Chordata" rank="species" species="rex">rex</taxonomicName>
|
||
</emphasis>
|
||
.
|
||
</emphasis>
|
||
The rows are organized by node, followed by the sequence of number of synontomorphies and individual variation by number, rank, and midranks. Boldfaced columns indicate values used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="77" pageNumber="78">
|
||
<table box="[516,1524,378,1628]" gridcols="7" gridrows="32" pageId="77" pageNumber="78">
|
||
<tr box="[516,1524,378,459]" gridrow="0" pageId="77" pageNumber="78">
|
||
<th box="[516,569,378,459]" gridcol="0" gridrow="0" pageId="77" pageNumber="78">Node</th>
|
||
<th box="[588,791,378,459]" gridcol="1" gridrow="0" pageId="77" pageNumber="78"># Synontomoprhies/ node</th>
|
||
<th box="[810,971,378,459]" gridcol="2" gridrow="0" pageId="77" pageNumber="78">Synontomorphy rank</th>
|
||
<th box="[992,1153,378,459]" gridcol="3" gridrow="0" pageId="77" pageNumber="78">Synontomorphy midrank</th>
|
||
<th box="[1173,1277,378,459]" gridcol="4" gridrow="0" pageId="77" pageNumber="78">Individual variation/ branch</th>
|
||
<th box="[1297,1401,378,459]" gridcol="5" gridrow="0" pageId="77" pageNumber="78">Individual variation rank</th>
|
||
<th box="[1420,1524,378,459]" gridcol="6" gridrow="0" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1524,378,400]" pageId="77" pageNumber="78">Individual</emphasis>
|
||
<emphasis bold="true" box="[1420,1510,407,459]" pageId="77" pageNumber="78">variation midrank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1524,478,499]" gridrow="1" pageId="77" pageNumber="78">
|
||
<th box="[516,569,478,499]" gridcol="0" gridrow="1" pageId="77" pageNumber="78">1</th>
|
||
<td box="[588,791,478,499]" gridcol="1" gridrow="1" pageId="77" pageNumber="78">0</td>
|
||
<td box="[810,971,478,499]" gridcol="2" gridrow="1" pageId="77" pageNumber="78">1</td>
|
||
<td box="[992,1153,478,499]" gridcol="3" gridrow="1" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,478,499]" pageId="77" pageNumber="78">1.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,478,499]" gridcol="4" gridrow="1" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,478,499]" gridcol="5" gridrow="1" pageId="77" pageNumber="78">1</td>
|
||
<td box="[1420,1524,478,499]" gridcol="6" gridrow="1" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,478,499]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,515,537]" gridrow="2" pageId="77" pageNumber="78">
|
||
<th box="[516,569,515,537]" gridcol="0" gridrow="2" pageId="77" pageNumber="78">2</th>
|
||
<td box="[588,791,515,537]" gridcol="1" gridrow="2" pageId="77" pageNumber="78">2</td>
|
||
<td box="[810,971,515,537]" gridcol="2" gridrow="2" pageId="77" pageNumber="78">4</td>
|
||
<td box="[992,1153,515,537]" gridcol="3" gridrow="2" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,516,537]" pageId="77" pageNumber="78">5.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,515,537]" gridcol="4" gridrow="2" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,515,537]" gridcol="5" gridrow="2" pageId="77" pageNumber="78">2</td>
|
||
<td box="[1420,1524,515,537]" gridcol="6" gridrow="2" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,515,536]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,553,574]" gridrow="3" pageId="77" pageNumber="78">
|
||
<th box="[516,569,553,574]" gridcol="0" gridrow="3" pageId="77" pageNumber="78">3</th>
|
||
<td box="[588,791,553,574]" gridcol="1" gridrow="3" pageId="77" pageNumber="78">0</td>
|
||
<td box="[810,971,553,574]" gridcol="2" gridrow="3" pageId="77" pageNumber="78">2</td>
|
||
<td box="[992,1153,553,574]" gridcol="3" gridrow="3" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,553,574]" pageId="77" pageNumber="78">1.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,553,574]" gridcol="4" gridrow="3" pageId="77" pageNumber="78">3</td>
|
||
<td box="[1297,1401,553,574]" gridcol="5" gridrow="3" pageId="77" pageNumber="78">11</td>
|
||
<td box="[1420,1524,553,574]" gridcol="6" gridrow="3" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,553,574]" pageId="77" pageNumber="78">12</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,591,612]" gridrow="4" pageId="77" pageNumber="78">
|
||
<th box="[516,569,591,612]" gridcol="0" gridrow="4" pageId="77" pageNumber="78">4</th>
|
||
<td box="[588,791,591,612]" gridcol="1" gridrow="4" pageId="77" pageNumber="78">3</td>
|
||
<td box="[810,971,591,612]" gridcol="2" gridrow="4" pageId="77" pageNumber="78">8</td>
|
||
<td box="[992,1153,591,612]" gridcol="3" gridrow="4" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,591,612]" pageId="77" pageNumber="78">8.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,591,612]" gridcol="4" gridrow="4" pageId="77" pageNumber="78">3</td>
|
||
<td box="[1297,1401,591,612]" gridcol="5" gridrow="4" pageId="77" pageNumber="78">12</td>
|
||
<td box="[1420,1524,591,612]" gridcol="6" gridrow="4" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,591,612]" pageId="77" pageNumber="78">12</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,628,650]" gridrow="5" pageId="77" pageNumber="78">
|
||
<th box="[516,569,628,650]" gridcol="0" gridrow="5" pageId="77" pageNumber="78">5</th>
|
||
<td box="[588,791,628,650]" gridcol="1" gridrow="5" pageId="77" pageNumber="78">50</td>
|
||
<td box="[810,971,628,650]" gridcol="2" gridrow="5" pageId="77" pageNumber="78">29</td>
|
||
<td box="[992,1153,628,650]" gridcol="3" gridrow="5" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,628,649]" pageId="77" pageNumber="78">29</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,628,650]" gridcol="4" gridrow="5" pageId="77" pageNumber="78">16</td>
|
||
<td box="[1297,1401,628,650]" gridcol="5" gridrow="5" pageId="77" pageNumber="78">26</td>
|
||
<td box="[1420,1524,628,650]" gridcol="6" gridrow="5" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,628,649]" pageId="77" pageNumber="78">26</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,666,688]" gridrow="6" pageId="77" pageNumber="78">
|
||
<th box="[516,569,666,688]" gridcol="0" gridrow="6" pageId="77" pageNumber="78">6</th>
|
||
<td box="[588,791,666,688]" gridcol="1" gridrow="6" pageId="77" pageNumber="78">90</td>
|
||
<td box="[810,971,666,688]" gridcol="2" gridrow="6" pageId="77" pageNumber="78">30</td>
|
||
<td box="[992,1153,666,688]" gridcol="3" gridrow="6" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,666,687]" pageId="77" pageNumber="78">30.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,666,688]" gridcol="4" gridrow="6" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,666,688]" gridcol="5" gridrow="6" pageId="77" pageNumber="78">3</td>
|
||
<td box="[1420,1524,666,688]" gridcol="6" gridrow="6" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,666,687]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,703,726]" gridrow="7" pageId="77" pageNumber="78">
|
||
<th box="[516,569,703,726]" gridcol="0" gridrow="7" pageId="77" pageNumber="78">6</th>
|
||
<td box="[588,791,703,726]" gridcol="1" gridrow="7" pageId="77" pageNumber="78">90</td>
|
||
<td box="[810,971,703,726]" gridcol="2" gridrow="7" pageId="77" pageNumber="78">31</td>
|
||
<td box="[992,1153,703,726]" gridcol="3" gridrow="7" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,703,724]" pageId="77" pageNumber="78">30.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,703,726]" gridcol="4" gridrow="7" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,703,726]" gridcol="5" gridrow="7" pageId="77" pageNumber="78">4</td>
|
||
<td box="[1420,1524,703,726]" gridcol="6" gridrow="7" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,703,724]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,741,763]" gridrow="8" pageId="77" pageNumber="78">
|
||
<th box="[516,569,741,763]" gridcol="0" gridrow="8" pageId="77" pageNumber="78">7</th>
|
||
<td box="[588,791,741,763]" gridcol="1" gridrow="8" pageId="77" pageNumber="78">16</td>
|
||
<td box="[810,971,741,763]" gridcol="2" gridrow="8" pageId="77" pageNumber="78">23</td>
|
||
<td box="[992,1153,741,763]" gridcol="3" gridrow="8" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,741,762]" pageId="77" pageNumber="78">23.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,741,763]" gridcol="4" gridrow="8" pageId="77" pageNumber="78">6</td>
|
||
<td box="[1297,1401,741,763]" gridcol="5" gridrow="8" pageId="77" pageNumber="78">18</td>
|
||
<td box="[1420,1524,741,763]" gridcol="6" gridrow="8" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,741,762]" pageId="77" pageNumber="78">18</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,779,800]" gridrow="9" pageId="77" pageNumber="78">
|
||
<th box="[516,569,779,800]" gridcol="0" gridrow="9" pageId="77" pageNumber="78">8</th>
|
||
<td box="[588,791,779,800]" gridcol="1" gridrow="9" pageId="77" pageNumber="78">31</td>
|
||
<td box="[810,971,779,800]" gridcol="2" gridrow="9" pageId="77" pageNumber="78">28</td>
|
||
<td box="[992,1153,779,800]" gridcol="3" gridrow="9" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,779,800]" pageId="77" pageNumber="78">28</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,779,800]" gridcol="4" gridrow="9" pageId="77" pageNumber="78">49</td>
|
||
<td box="[1297,1401,779,800]" gridcol="5" gridrow="9" pageId="77" pageNumber="78">31</td>
|
||
<td box="[1420,1524,779,800]" gridcol="6" gridrow="9" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,779,800]" pageId="77" pageNumber="78">31</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,816,838]" gridrow="10" pageId="77" pageNumber="78">
|
||
<th box="[516,569,816,838]" gridcol="0" gridrow="10" pageId="77" pageNumber="78">9</th>
|
||
<td box="[588,791,816,838]" gridcol="1" gridrow="10" pageId="77" pageNumber="78">16</td>
|
||
<td box="[810,971,816,838]" gridcol="2" gridrow="10" pageId="77" pageNumber="78">24</td>
|
||
<td box="[992,1153,816,838]" gridcol="3" gridrow="10" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,816,837]" pageId="77" pageNumber="78">23.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,816,838]" gridcol="4" gridrow="10" pageId="77" pageNumber="78">6</td>
|
||
<td box="[1297,1401,816,838]" gridcol="5" gridrow="10" pageId="77" pageNumber="78">18</td>
|
||
<td box="[1420,1524,816,838]" gridcol="6" gridrow="10" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,816,837]" pageId="77" pageNumber="78">18</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,854,875]" gridrow="11" pageId="77" pageNumber="78">
|
||
<th box="[516,569,854,875]" gridcol="0" gridrow="11" pageId="77" pageNumber="78">10</th>
|
||
<td box="[588,791,854,875]" gridcol="1" gridrow="11" pageId="77" pageNumber="78">12</td>
|
||
<td box="[810,971,854,875]" gridcol="2" gridrow="11" pageId="77" pageNumber="78">19</td>
|
||
<td box="[992,1153,854,875]" gridcol="3" gridrow="11" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,854,875]" pageId="77" pageNumber="78">19</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,854,875]" gridcol="4" gridrow="11" pageId="77" pageNumber="78">42</td>
|
||
<td box="[1297,1401,854,875]" gridcol="5" gridrow="11" pageId="77" pageNumber="78">29</td>
|
||
<td box="[1420,1524,854,875]" gridcol="6" gridrow="11" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,854,875]" pageId="77" pageNumber="78">29</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,891,913]" gridrow="12" pageId="77" pageNumber="78">
|
||
<th box="[516,569,891,913]" gridcol="0" gridrow="12" pageId="77" pageNumber="78">11</th>
|
||
<td box="[588,791,891,913]" gridcol="1" gridrow="12" pageId="77" pageNumber="78">9</td>
|
||
<td box="[810,971,891,913]" gridcol="2" gridrow="12" pageId="77" pageNumber="78">14</td>
|
||
<td box="[992,1153,891,913]" gridcol="3" gridrow="12" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,891,912]" pageId="77" pageNumber="78">15</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,891,913]" gridcol="4" gridrow="12" pageId="77" pageNumber="78">17</td>
|
||
<td box="[1297,1401,891,913]" gridcol="5" gridrow="12" pageId="77" pageNumber="78">27</td>
|
||
<td box="[1420,1524,891,913]" gridcol="6" gridrow="12" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,891,912]" pageId="77" pageNumber="78">27</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,929,951]" gridrow="13" pageId="77" pageNumber="78">
|
||
<th box="[516,569,929,951]" gridcol="0" gridrow="13" pageId="77" pageNumber="78">12</th>
|
||
<td box="[588,791,929,951]" gridcol="1" gridrow="13" pageId="77" pageNumber="78">9</td>
|
||
<td box="[810,971,929,951]" gridcol="2" gridrow="13" pageId="77" pageNumber="78">15</td>
|
||
<td box="[992,1153,929,951]" gridcol="3" gridrow="13" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,929,950]" pageId="77" pageNumber="78">15</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,929,951]" gridcol="4" gridrow="13" pageId="77" pageNumber="78">2</td>
|
||
<td box="[1297,1401,929,951]" gridcol="5" gridrow="13" pageId="77" pageNumber="78">10</td>
|
||
<td box="[1420,1524,929,951]" gridcol="6" gridrow="13" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,929,950]" pageId="77" pageNumber="78">10</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,966,989]" gridrow="14" pageId="77" pageNumber="78">
|
||
<th box="[516,569,966,989]" gridcol="0" gridrow="14" pageId="77" pageNumber="78">12</th>
|
||
<td box="[588,791,966,989]" gridcol="1" gridrow="14" pageId="77" pageNumber="78">9</td>
|
||
<td box="[810,971,966,989]" gridcol="2" gridrow="14" pageId="77" pageNumber="78">16</td>
|
||
<td box="[992,1153,966,989]" gridcol="3" gridrow="14" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,966,987]" pageId="77" pageNumber="78">15</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,966,989]" gridcol="4" gridrow="14" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,966,989]" gridcol="5" gridrow="14" pageId="77" pageNumber="78">5</td>
|
||
<td box="[1420,1524,966,989]" gridcol="6" gridrow="14" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,966,987]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1004,1026]" gridrow="15" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1004,1026]" gridcol="0" gridrow="15" pageId="77" pageNumber="78">13</th>
|
||
<td box="[588,791,1004,1026]" gridcol="1" gridrow="15" pageId="77" pageNumber="78">20</td>
|
||
<td box="[810,971,1004,1026]" gridcol="2" gridrow="15" pageId="77" pageNumber="78">26</td>
|
||
<td box="[992,1153,1004,1026]" gridcol="3" gridrow="15" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1004,1025]" pageId="77" pageNumber="78">26.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1004,1026]" gridcol="4" gridrow="15" pageId="77" pageNumber="78">5</td>
|
||
<td box="[1297,1401,1004,1026]" gridcol="5" gridrow="15" pageId="77" pageNumber="78">15</td>
|
||
<td box="[1420,1524,1004,1026]" gridcol="6" gridrow="15" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1462,1004,1025]" pageId="77" pageNumber="78">15.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1042,1063]" gridrow="16" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1042,1063]" gridcol="0" gridrow="16" pageId="77" pageNumber="78">13</th>
|
||
<td box="[588,791,1042,1063]" gridcol="1" gridrow="16" pageId="77" pageNumber="78">20</td>
|
||
<td box="[810,971,1042,1063]" gridcol="2" gridrow="16" pageId="77" pageNumber="78">27</td>
|
||
<td box="[992,1153,1042,1063]" gridcol="3" gridrow="16" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1042,1063]" pageId="77" pageNumber="78">26.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1042,1063]" gridcol="4" gridrow="16" pageId="77" pageNumber="78">9</td>
|
||
<td box="[1297,1401,1042,1063]" gridcol="5" gridrow="16" pageId="77" pageNumber="78">21</td>
|
||
<td box="[1420,1524,1042,1063]" gridcol="6" gridrow="16" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1042,1063]" pageId="77" pageNumber="78">21</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1079,1101]" gridrow="17" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1079,1101]" gridcol="0" gridrow="17" pageId="77" pageNumber="78">14</th>
|
||
<td box="[588,791,1079,1101]" gridcol="1" gridrow="17" pageId="77" pageNumber="78">10</td>
|
||
<td box="[810,971,1079,1101]" gridcol="2" gridrow="17" pageId="77" pageNumber="78">17</td>
|
||
<td box="[992,1153,1079,1101]" gridcol="3" gridrow="17" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1079,1100]" pageId="77" pageNumber="78">17.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1079,1101]" gridcol="4" gridrow="17" pageId="77" pageNumber="78">10</td>
|
||
<td box="[1297,1401,1079,1101]" gridcol="5" gridrow="17" pageId="77" pageNumber="78">22</td>
|
||
<td box="[1420,1524,1079,1101]" gridcol="6" gridrow="17" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1079,1100]" pageId="77" pageNumber="78">22</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1117,1139]" gridrow="18" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1117,1139]" gridcol="0" gridrow="18" pageId="77" pageNumber="78">14</th>
|
||
<td box="[588,791,1117,1139]" gridcol="1" gridrow="18" pageId="77" pageNumber="78">10</td>
|
||
<td box="[810,971,1117,1139]" gridcol="2" gridrow="18" pageId="77" pageNumber="78">18</td>
|
||
<td box="[992,1153,1117,1139]" gridcol="3" gridrow="18" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1117,1138]" pageId="77" pageNumber="78">17.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1117,1139]" gridcol="4" gridrow="18" pageId="77" pageNumber="78">5</td>
|
||
<td box="[1297,1401,1117,1139]" gridcol="5" gridrow="18" pageId="77" pageNumber="78">16</td>
|
||
<td box="[1420,1524,1117,1139]" gridcol="6" gridrow="18" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1462,1117,1138]" pageId="77" pageNumber="78">15.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1155,1177]" gridrow="19" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1155,1177]" gridcol="0" gridrow="19" pageId="77" pageNumber="78">15</th>
|
||
<td box="[588,791,1155,1177]" gridcol="1" gridrow="19" pageId="77" pageNumber="78">3</td>
|
||
<td box="[810,971,1155,1177]" gridcol="2" gridrow="19" pageId="77" pageNumber="78">9</td>
|
||
<td box="[992,1153,1155,1177]" gridcol="3" gridrow="19" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,1155,1176]" pageId="77" pageNumber="78">8.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1155,1177]" gridcol="4" gridrow="19" pageId="77" pageNumber="78">3</td>
|
||
<td box="[1297,1401,1155,1177]" gridcol="5" gridrow="19" pageId="77" pageNumber="78">13</td>
|
||
<td box="[1420,1524,1155,1177]" gridcol="6" gridrow="19" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1155,1176]" pageId="77" pageNumber="78">12</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1192,1214]" gridrow="20" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1192,1214]" gridcol="0" gridrow="20" pageId="77" pageNumber="78">16</th>
|
||
<td box="[588,791,1192,1214]" gridcol="1" gridrow="20" pageId="77" pageNumber="78">1</td>
|
||
<td box="[810,971,1192,1214]" gridcol="2" gridrow="20" pageId="77" pageNumber="78">3</td>
|
||
<td box="[992,1153,1192,1214]" gridcol="3" gridrow="20" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1004,1192,1213]" pageId="77" pageNumber="78">3</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1192,1214]" gridcol="4" gridrow="20" pageId="77" pageNumber="78">4</td>
|
||
<td box="[1297,1401,1192,1214]" gridcol="5" gridrow="20" pageId="77" pageNumber="78">14</td>
|
||
<td box="[1420,1524,1192,1214]" gridcol="6" gridrow="20" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1192,1213]" pageId="77" pageNumber="78">14</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1230,1252]" gridrow="21" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1230,1252]" gridcol="0" gridrow="21" pageId="77" pageNumber="78">17</th>
|
||
<td box="[588,791,1230,1252]" gridcol="1" gridrow="21" pageId="77" pageNumber="78">2</td>
|
||
<td box="[810,971,1230,1252]" gridcol="2" gridrow="21" pageId="77" pageNumber="78">5</td>
|
||
<td box="[992,1153,1230,1252]" gridcol="3" gridrow="21" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,1230,1251]" pageId="77" pageNumber="78">5.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1230,1252]" gridcol="4" gridrow="21" pageId="77" pageNumber="78">15</td>
|
||
<td box="[1297,1401,1230,1252]" gridcol="5" gridrow="21" pageId="77" pageNumber="78">25</td>
|
||
<td box="[1420,1524,1230,1252]" gridcol="6" gridrow="21" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1230,1251]" pageId="77" pageNumber="78">25</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1267,1289]" gridrow="22" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1267,1289]" gridcol="0" gridrow="22" pageId="77" pageNumber="78">17</th>
|
||
<td box="[588,791,1267,1289]" gridcol="1" gridrow="22" pageId="77" pageNumber="78">2</td>
|
||
<td box="[810,971,1267,1289]" gridcol="2" gridrow="22" pageId="77" pageNumber="78">6</td>
|
||
<td box="[992,1153,1267,1289]" gridcol="3" gridrow="22" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,1268,1289]" pageId="77" pageNumber="78">5.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1267,1289]" gridcol="4" gridrow="22" pageId="77" pageNumber="78">41</td>
|
||
<td box="[1297,1401,1267,1289]" gridcol="5" gridrow="22" pageId="77" pageNumber="78">28</td>
|
||
<td box="[1420,1524,1267,1289]" gridcol="6" gridrow="22" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1267,1288]" pageId="77" pageNumber="78">28</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1305,1326]" gridrow="23" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1305,1326]" gridcol="0" gridrow="23" pageId="77" pageNumber="78">17</th>
|
||
<td box="[588,791,1305,1326]" gridcol="1" gridrow="23" pageId="77" pageNumber="78">2</td>
|
||
<td box="[810,971,1305,1326]" gridcol="2" gridrow="23" pageId="77" pageNumber="78">7</td>
|
||
<td box="[992,1153,1305,1326]" gridcol="3" gridrow="23" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1022,1305,1326]" pageId="77" pageNumber="78">5.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1305,1326]" gridcol="4" gridrow="23" pageId="77" pageNumber="78">47</td>
|
||
<td box="[1297,1401,1305,1326]" gridcol="5" gridrow="23" pageId="77" pageNumber="78">30</td>
|
||
<td box="[1420,1524,1305,1326]" gridcol="6" gridrow="23" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1305,1326]" pageId="77" pageNumber="78">30</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1342,1365]" gridrow="24" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1342,1365]" gridcol="0" gridrow="24" pageId="77" pageNumber="78">18</th>
|
||
<td box="[588,791,1342,1365]" gridcol="1" gridrow="24" pageId="77" pageNumber="78">6</td>
|
||
<td box="[810,971,1342,1365]" gridcol="2" gridrow="24" pageId="77" pageNumber="78">10</td>
|
||
<td box="[992,1153,1342,1365]" gridcol="3" gridrow="24" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1342,1363]" pageId="77" pageNumber="78">10.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1342,1365]" gridcol="4" gridrow="24" pageId="77" pageNumber="78">12</td>
|
||
<td box="[1297,1401,1342,1365]" gridcol="5" gridrow="24" pageId="77" pageNumber="78">24</td>
|
||
<td box="[1420,1524,1342,1365]" gridcol="6" gridrow="24" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1342,1363]" pageId="77" pageNumber="78">24</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1380,1402]" gridrow="25" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1380,1402]" gridcol="0" gridrow="25" pageId="77" pageNumber="78">18</th>
|
||
<td box="[588,791,1380,1402]" gridcol="1" gridrow="25" pageId="77" pageNumber="78">6</td>
|
||
<td box="[810,971,1380,1402]" gridcol="2" gridrow="25" pageId="77" pageNumber="78">11</td>
|
||
<td box="[992,1153,1380,1402]" gridcol="3" gridrow="25" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1380,1401]" pageId="77" pageNumber="78">10.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1380,1402]" gridcol="4" gridrow="25" pageId="77" pageNumber="78">8</td>
|
||
<td box="[1297,1401,1380,1402]" gridcol="5" gridrow="25" pageId="77" pageNumber="78">20</td>
|
||
<td box="[1420,1524,1380,1402]" gridcol="6" gridrow="25" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1380,1401]" pageId="77" pageNumber="78">20</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1418,1440]" gridrow="26" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1418,1440]" gridcol="0" gridrow="26" pageId="77" pageNumber="78">19</th>
|
||
<td box="[588,791,1418,1440]" gridcol="1" gridrow="26" pageId="77" pageNumber="78">15</td>
|
||
<td box="[810,971,1418,1440]" gridcol="2" gridrow="26" pageId="77" pageNumber="78">20</td>
|
||
<td box="[992,1153,1418,1440]" gridcol="3" gridrow="26" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,1418,1439]" pageId="77" pageNumber="78">21</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1418,1440]" gridcol="4" gridrow="26" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,1418,1440]" gridcol="5" gridrow="26" pageId="77" pageNumber="78">6</td>
|
||
<td box="[1420,1524,1418,1440]" gridcol="6" gridrow="26" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,1418,1439]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1455,1477]" gridrow="27" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1455,1477]" gridcol="0" gridrow="27" pageId="77" pageNumber="78">19</th>
|
||
<td box="[588,791,1455,1477]" gridcol="1" gridrow="27" pageId="77" pageNumber="78">15</td>
|
||
<td box="[810,971,1455,1477]" gridcol="2" gridrow="27" pageId="77" pageNumber="78">21</td>
|
||
<td box="[992,1153,1455,1477]" gridcol="3" gridrow="27" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,1455,1476]" pageId="77" pageNumber="78">21</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1455,1477]" gridcol="4" gridrow="27" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,1455,1477]" gridcol="5" gridrow="27" pageId="77" pageNumber="78">7</td>
|
||
<td box="[1420,1524,1455,1477]" gridcol="6" gridrow="27" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,1455,1476]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1493,1515]" gridrow="28" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1493,1515]" gridcol="0" gridrow="28" pageId="77" pageNumber="78">19</th>
|
||
<td box="[588,791,1493,1515]" gridcol="1" gridrow="28" pageId="77" pageNumber="78">15</td>
|
||
<td box="[810,971,1493,1515]" gridcol="2" gridrow="28" pageId="77" pageNumber="78">22</td>
|
||
<td box="[992,1153,1493,1515]" gridcol="3" gridrow="28" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,1493,1514]" pageId="77" pageNumber="78">21</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1493,1515]" gridcol="4" gridrow="28" pageId="77" pageNumber="78">1</td>
|
||
<td box="[1297,1401,1493,1515]" gridcol="5" gridrow="28" pageId="77" pageNumber="78">9</td>
|
||
<td box="[1420,1524,1493,1515]" gridcol="6" gridrow="28" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1432,1493,1514]" pageId="77" pageNumber="78">9</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1530,1552]" gridrow="29" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1530,1552]" gridcol="0" gridrow="29" pageId="77" pageNumber="78">20</th>
|
||
<td box="[588,791,1530,1552]" gridcol="1" gridrow="29" pageId="77" pageNumber="78">7</td>
|
||
<td box="[810,971,1530,1552]" gridcol="2" gridrow="29" pageId="77" pageNumber="78">12</td>
|
||
<td box="[992,1153,1530,1552]" gridcol="3" gridrow="29" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1530,1551]" pageId="77" pageNumber="78">12.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1530,1552]" gridcol="4" gridrow="29" pageId="77" pageNumber="78">11</td>
|
||
<td box="[1297,1401,1530,1552]" gridcol="5" gridrow="29" pageId="77" pageNumber="78">23</td>
|
||
<td box="[1420,1524,1530,1552]" gridcol="6" gridrow="29" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1530,1551]" pageId="77" pageNumber="78">23</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1568,1590]" gridrow="30" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1568,1590]" gridcol="0" gridrow="30" pageId="77" pageNumber="78">20</th>
|
||
<td box="[588,791,1568,1590]" gridcol="1" gridrow="30" pageId="77" pageNumber="78">7</td>
|
||
<td box="[810,971,1568,1590]" gridcol="2" gridrow="30" pageId="77" pageNumber="78">13</td>
|
||
<td box="[992,1153,1568,1590]" gridcol="3" gridrow="30" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1034,1568,1589]" pageId="77" pageNumber="78">12.5</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1568,1590]" gridcol="4" gridrow="30" pageId="77" pageNumber="78">6</td>
|
||
<td box="[1297,1401,1568,1590]" gridcol="5" gridrow="30" pageId="77" pageNumber="78">18</td>
|
||
<td box="[1420,1524,1568,1590]" gridcol="6" gridrow="30" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1444,1568,1589]" pageId="77" pageNumber="78">18</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1524,1606,1628]" gridrow="31" pageId="77" pageNumber="78">
|
||
<th box="[516,569,1606,1628]" gridcol="0" gridrow="31" pageId="77" pageNumber="78">21</th>
|
||
<td box="[588,791,1606,1628]" gridcol="1" gridrow="31" pageId="77" pageNumber="78">19</td>
|
||
<td box="[810,971,1606,1628]" gridcol="2" gridrow="31" pageId="77" pageNumber="78">25</td>
|
||
<td box="[992,1153,1606,1628]" gridcol="3" gridrow="31" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[992,1016,1606,1627]" pageId="77" pageNumber="78">25</emphasis>
|
||
</td>
|
||
<td box="[1173,1277,1606,1628]" gridcol="4" gridrow="31" pageId="77" pageNumber="78">0</td>
|
||
<td box="[1297,1401,1606,1628]" gridcol="5" gridrow="31" pageId="77" pageNumber="78">8</td>
|
||
<td box="[1420,1524,1606,1628]" gridcol="6" gridrow="31" pageId="77" pageNumber="78">
|
||
<emphasis bold="true" box="[1420,1450,1606,1627]" pageId="77" pageNumber="78">4.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825038" ID-Zenodo-Dep="5825038" httpUri="https://zenodo.org/record/5825038/files/figure.png" pageId="78" pageNumber="79" startId="78.[524,589,656,678]" targetBox="[500,1298,234,623]" targetPageId="78">
|
||
<paragraph blockId="78.[524,1515,656,854]" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" pageId="78" pageNumber="79">
|
||
Figure 32 Bivariate scatterplot showing the congruence between individual variation per specimen per node compared with the number of unambiguously optimized synontomorphies per node in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,713,715,736]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="78" pageNumber="79" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[524,713,715,736]" italics="true" pageId="78" pageNumber="79">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The number of synontomorphies per node are along the
|
||
<emphasis box="[1302,1313,715,736]" italics="true" pageId="78" pageNumber="79">x</emphasis>
|
||
-axis; the amount of individual variation (i.e., unambiguously optimized character states per branch) is along the
|
||
<emphasis box="[1403,1413,745,766]" italics="true" pageId="78" pageNumber="79">y</emphasis>
|
||
-axis. Both values increase away from the origin. If the amount of individual variation is controlled by the number of synontomorphies per node, then the variables should increase monotonically. In this case, no congruence is seen between the variables in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[834,890,832,853]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="78" pageNumber="79" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[834,890,832,853]" italics="true" pageId="78" pageNumber="79">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Full-size DOI: 10.7717/peerj.9192/fig-32
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-Table-UUID="DF5C84E2FF8AFFE6FDEBFC1734C8FBB0" httpUri="http://table.plazi.org/id/DF5C84E2FF8AFFE6FDEBFC1734C8FBB0" pageId="78" pageNumber="79" startId="78.[525,580,927,948]" targetBox="[516,1525,1056,1366]" targetIsTable="true" targetPageId="78">
|
||
<paragraph blockId="78.[525,1516,926,1036]" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" pageId="78" pageNumber="79">Table 21 Summary of the number of synontomorphies and individual variation per craniomandibular module.</emphasis>
|
||
Raw and corrected data used in the correlation test between the number of synontomorphies in each skull module with the amount of individual variation per module. Boldface indicates the ranks used in the correlation test.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="78" pageNumber="79">
|
||
<table box="[516,1525,1056,1366]" gridcols="6" gridrows="7" pageId="78" pageNumber="79">
|
||
<tr box="[516,1525,1056,1137]" gridrow="0" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1056,1137]" gridcol="0" gridrow="0" pageId="78" pageNumber="79">Module</th>
|
||
<th box="[675,851,1056,1137]" gridcol="1" gridrow="0" pageId="78" pageNumber="79">Synontomorphies</th>
|
||
<th box="[909,1085,1056,1137]" gridcol="2" gridrow="0" pageId="78" pageNumber="79">Synontomorphies rank</th>
|
||
<th box="[1144,1248,1056,1137]" gridcol="3" gridrow="0" pageId="78" pageNumber="79">Individual variation</th>
|
||
<th box="[1282,1386,1056,1137]" gridcol="4" gridrow="0" pageId="78" pageNumber="79">Individual variation rank</th>
|
||
<th box="[1421,1525,1056,1137]" gridcol="5" gridrow="0" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1525,1056,1078]" pageId="78" pageNumber="79">Individual</emphasis>
|
||
<emphasis bold="true" box="[1421,1511,1086,1108]" pageId="78" pageNumber="79">variation</emphasis>
|
||
<emphasis bold="true" box="[1421,1507,1115,1137]" pageId="78" pageNumber="79">midrank</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1525,1156,1178]" gridrow="1" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1156,1178]" gridcol="0" gridrow="1" pageId="78" pageNumber="79">Skull roof</th>
|
||
<td box="[675,851,1156,1178]" gridcol="1" gridrow="1" pageId="78" pageNumber="79">117</td>
|
||
<td box="[909,1085,1156,1178]" gridcol="2" gridrow="1" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1156,1177]" pageId="78" pageNumber="79">1</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1156,1178]" gridcol="3" gridrow="1" pageId="78" pageNumber="79">100</td>
|
||
<td box="[1282,1386,1156,1178]" gridcol="4" gridrow="1" pageId="78" pageNumber="79">1</td>
|
||
<td box="[1421,1525,1156,1178]" gridcol="5" gridrow="1" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1433,1156,1177]" pageId="78" pageNumber="79">1</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1194,1216]" gridrow="2" pageId="78" pageNumber="79">
|
||
<th box="[516,851,1194,1216]" colspan="2" colspanRight="1" gridcol="0" gridrow="2" pageId="78" pageNumber="79">Snout & palate 60</th>
|
||
<td box="[909,1085,1194,1216]" gridcol="2" gridrow="2" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1194,1215]" pageId="78" pageNumber="79">2</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1194,1216]" gridcol="3" gridrow="2" pageId="78" pageNumber="79">57</td>
|
||
<td box="[1282,1386,1194,1216]" gridcol="4" gridrow="2" pageId="78" pageNumber="79">2</td>
|
||
<td box="[1421,1525,1194,1216]" gridcol="5" gridrow="2" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1433,1194,1215]" pageId="78" pageNumber="79">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1231,1253]" gridrow="3" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1231,1253]" gridcol="0" gridrow="3" pageId="78" pageNumber="79">Parietal</th>
|
||
<td box="[675,851,1231,1253]" gridcol="1" gridrow="3" pageId="78" pageNumber="79">12</td>
|
||
<td box="[909,1085,1231,1253]" gridcol="2" gridrow="3" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1231,1252]" pageId="78" pageNumber="79">6</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1231,1253]" gridcol="3" gridrow="3" pageId="78" pageNumber="79">23</td>
|
||
<td box="[1282,1386,1231,1253]" gridcol="4" gridrow="3" pageId="78" pageNumber="79">5</td>
|
||
<td box="[1421,1525,1231,1253]" gridcol="5" gridrow="3" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1451,1232,1253]" pageId="78" pageNumber="79">5.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1269,1291]" gridrow="4" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1269,1291]" gridcol="0" gridrow="4" pageId="78" pageNumber="79">Suspensorium</th>
|
||
<td box="[675,851,1269,1291]" gridcol="1" gridrow="4" pageId="78" pageNumber="79">13</td>
|
||
<td box="[909,1085,1269,1291]" gridcol="2" gridrow="4" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1269,1290]" pageId="78" pageNumber="79">5</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1269,1291]" gridcol="3" gridrow="4" pageId="78" pageNumber="79">23</td>
|
||
<td box="[1282,1386,1269,1291]" gridcol="4" gridrow="4" pageId="78" pageNumber="79">6</td>
|
||
<td box="[1421,1525,1269,1291]" gridcol="5" gridrow="4" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1451,1269,1290]" pageId="78" pageNumber="79">5.5</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1307,1329]" gridrow="5" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1307,1329]" gridcol="0" gridrow="5" pageId="78" pageNumber="79">Braincase</th>
|
||
<td box="[675,851,1307,1329]" gridcol="1" gridrow="5" pageId="78" pageNumber="79">25</td>
|
||
<td box="[909,1085,1307,1329]" gridcol="2" gridrow="5" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1307,1328]" pageId="78" pageNumber="79">4</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1307,1329]" gridcol="3" gridrow="5" pageId="78" pageNumber="79">29</td>
|
||
<td box="[1282,1386,1307,1329]" gridcol="4" gridrow="5" pageId="78" pageNumber="79">3</td>
|
||
<td box="[1421,1525,1307,1329]" gridcol="5" gridrow="5" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1433,1307,1328]" pageId="78" pageNumber="79">3</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1525,1344,1366]" gridrow="6" pageId="78" pageNumber="79">
|
||
<th box="[516,651,1344,1366]" gridcol="0" gridrow="6" pageId="78" pageNumber="79">Mandible</th>
|
||
<td box="[675,851,1344,1366]" gridcol="1" gridrow="6" pageId="78" pageNumber="79">27</td>
|
||
<td box="[909,1085,1344,1366]" gridcol="2" gridrow="6" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[909,921,1344,1365]" pageId="78" pageNumber="79">3</emphasis>
|
||
</td>
|
||
<td box="[1144,1248,1344,1366]" gridcol="3" gridrow="6" pageId="78" pageNumber="79">27</td>
|
||
<td box="[1282,1386,1344,1366]" gridcol="4" gridrow="6" pageId="78" pageNumber="79">4</td>
|
||
<td box="[1421,1525,1344,1366]" gridcol="5" gridrow="6" pageId="78" pageNumber="79">
|
||
<emphasis bold="true" box="[1421,1433,1344,1365]" pageId="78" pageNumber="79">4</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="78.[498,1542,1426,1931]" pageId="78" pageNumber="79">
|
||
Among all the characters that are optimized unambiguously as individual variation, the number of reversals (from a mature to an immature character state) is close to the number of progressions (from an immature state to the mature condition), 165
|
||
<emphasis box="[1434,1449,1627,1652]" italics="true" pageId="78" pageNumber="79">–</emphasis>
|
||
175, respectively (
|
||
<figureCitation box="[653,718,1666,1692]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="78" pageNumber="79">Fig. 2</figureCitation>
|
||
; Data S5). Progressions are the only type of change in the first seven growth stages; after that, progressions outnumber reversals until the 11th (exemplar:
|
||
<materialsCitation box="[498,734,1745,1772]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="78" pageNumber="79" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
) whereupon reversals establish their numerically dominant and constant presence (
|
||
<figureCitation box="[726,792,1785,1811]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="78" pageNumber="79">Fig. 2</figureCitation>
|
||
; Data S5).
|
||
</paragraph>
|
||
<paragraph blockId="78.[498,1542,1426,1931]" lastBlockId="79.[498,1542,234,858]" lastPageId="79" lastPageNumber="80" pageId="78" pageNumber="79">Only eight of the 165 reversals pertain to characters of display features, which include the presence of the ridge around the external antorbital fossa of the maxilla, the texture of the dorsal ramus of the lacrimal, and the height and size of the cornual process of the postorbital (Data S5). The low frequency of variation in cephalic ornaments indicates that these presumed intraspecific display structures are not any more labile than nonornamental characteristics.</paragraph>
|
||
<paragraph blockId="79.[498,1542,234,858]" pageId="79" pageNumber="80">
|
||
In summary, individual variation is seen across the entire skeleton, although it is more frequent in the skull and jaws (274) than in the postcranium (26). Almost certainly this discrepancy is, in part, an artifact of the lower number of characters scored for postcranial bones. Individual variation is not limited to any single domain; for example, in
|
||
<materialsCitation box="[530,706,513,540]" collectionCode="LACM" pageId="79" pageNumber="80" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
, a specimen with 33 optimizations, cranial variation includes pneumatization, bone shape, muscle scar orientation, alveolus size, foramen magnum size, skull frame, neurovascular foramen position, and alveolar skirt presence, to name a few. In the same specimen, postcranial variation includes, among others, the form of the scapular shaft, prominence of a scapular muscle scar, size of the cuppedicus fossa, form of the ischium, form of the distal joint surface on the fibula, and the orientation of the dorsal margin of the medial fossa of the fibula. As such, individual variation has an unlimited presence across the entire skeleton, which limits the precision of estimating relative maturity without a comprehensive character analysis such as the one presented here.
|
||
</paragraph>
|
||
<paragraph blockId="79.[498,1542,903,1932]" box="[498,804,903,933]" pageId="79" pageNumber="80">
|
||
<heading bold="true" box="[498,804,903,933]" fontSize="12" level="2" pageId="79" pageNumber="80" reason="0">
|
||
<emphasis bold="true" box="[498,804,903,933]" pageId="79" pageNumber="80">Oversplit characters</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="79.[498,1542,903,1932]" pageId="79" pageNumber="80">
|
||
A review of the synontomophies provides the opportunity for identifying characters that are oversplit (Data S4;
|
||
<tableCitation box="[824,925,989,1015]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="79" pageNumber="80" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). An oversplit character is defined here as two or more synontomorphies that support the same node and are subsets of the same morphological structure; for example, if the glenoid fossa of the scapula and coracoid are reoriented at the same node, then it is simplest to consider them to be the same change instead of two discrete changes. In contrast, if the glenoid fossa of the scapula changes orientation at an early node, whereas the glenoid fossa of the coracoid changes at a later node, then it is reasonable to treat them as independent characters.
|
||
</paragraph>
|
||
<paragraph blockId="79.[498,1542,903,1932]" pageId="79" pageNumber="80">
|
||
In growth stage 5, several characters might be oversplit, including several pertaining to skull height, the antorbital fossa, dorsal margin of the postorbital, facial subcutaneous texture, and maxillary tooth count (
|
||
<tableCitation box="[916,1016,1347,1373]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="79" pageNumber="80" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). In growth stage 6, characters of inflation of the dorsal ramus of the lacrimal and skull frame might be oversplit (
|
||
<tableCitation box="[1298,1398,1387,1413]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="79" pageNumber="80" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). In stage 7, oversplit characters pertain to inflation of the dorsal ramus of the lacrimal (
|
||
<tableCitation box="[1395,1496,1427,1453]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="79" pageNumber="80" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). In stage 9, oversplit characters might pertain to alveolus size (
|
||
<tableCitation box="[1223,1324,1467,1493]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="79" pageNumber="80" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). The test of these hypotheses is the addition of new specimens to the analysis that are sufficiently complete and of the appropriate growth stage. For instance, the great morphological differences between growth stages 4 and 5, and 5 and 6, indicate that specimens of intermediate morphology have not yet been found, and so the numerous changes seen in growth stages 5 and 6 that appear to be oversplit may have not developed simultaneously.
|
||
</paragraph>
|
||
<paragraph blockId="79.[498,1542,903,1932]" lastBlockId="80.[498,1542,1168,1712]" lastPageId="80" lastPageNumber="81" pageId="79" pageNumber="80">
|
||
Regardless, the effect of collapsing hypothetically overplit characters into a single transformation series was tested by running a second analysis. The transformation series were compared in three steps: (1) side-by-side comparison of possibly homologous transformation series, (2) matching transformation series were grouped together, (3) exemplar transformation series (i.e., those scored for the maximum number of specimens) were identified, and (4) the analysis was run with only the exemplar transformation series whereas the redundant transformation series were excluded from the analysis (
|
||
<tableCitation box="[608,710,1208,1234]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="80" pageNumber="81" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
). In cases where the transformation series are different, and so clearly test different morphologies or different aspects of the same structure, the characters were regarded as independent. Based on differences between transformation series, the characters pertaining to the antorbital fossa, dorsal margin of the postorbital, maxillary tooth count, lacrimal inflation, and alveolus size were not regarded as oversplit (
|
||
<tableCitation box="[1426,1525,1367,1394]" captionStart="Table 22" captionStartId="80.[117,172,248,269]" captionTargetBox="[108,1530,437,1093]" captionText="Table 22 Comparison of hypothetically oversplit characters in the data set for Tyrannosaurus rex ontogeny with decisions for deletion of redundant transformation series. The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not." httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="80" pageNumber="81" tableUuid="DF5C84E2FF94FFF8FF93FF703611FE1C">Table 22</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FF94FFF8FF93FF703611FE1C" httpUri="http://table.plazi.org/id/DF5C84E2FF94FFF8FF93FF703611FE1C" pageId="80" pageNumber="81" startId="80.[117,172,248,269]" targetBox="[108,1530,437,1093]" targetIsTable="true" targetPageId="80">
|
||
<paragraph blockId="80.[117,1516,248,416]" pageId="80" pageNumber="81">
|
||
<emphasis bold="true" pageId="80" pageNumber="81">
|
||
Table 22 Comparison of hypothetically oversplit characters in the data set for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[921,1108,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="80" pageNumber="81" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[921,1108,249,270]" italics="true" pageId="80" pageNumber="81">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
ontogeny with decisions for deletion of redundant transformation series.
|
||
</emphasis>
|
||
The rows are organized by growth stage, followed by the skull region, identification of oversplit characters, and the decisions (based on redundancy) to retain (=exemplar) or exclude transformation series from the analysis. Identical transformation series have matching codings, although one might be more complete than the other. Groups of identical transformation series are separated by semicolons. Exemplar transformation series are the most completely coded examples of their sets and were run in the analysis, whereas excluded transformations series were not.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="80" pageNumber="81">
|
||
<table box="[108,1530,437,1093]" gridcols="6" gridrows="10" pageId="80" pageNumber="81">
|
||
<tr box="[108,1530,437,517]" gridrow="0" pageId="80" pageNumber="81">
|
||
<th box="[108,184,437,517]" gridcol="0" gridrow="0" pageId="80" pageNumber="81">Growth stage</th>
|
||
<th box="[227,380,437,517]" gridcol="1" gridrow="0" pageId="80" pageNumber="81">Domain</th>
|
||
<th box="[422,709,437,517]" gridcol="2" gridrow="0" pageId="80" pageNumber="81">Hypothetically oversplit characters</th>
|
||
<th box="[744,1032,437,517]" gridcol="3" gridrow="0" pageId="80" pageNumber="81">Identical transformation series</th>
|
||
<th box="[1067,1282,437,517]" gridcol="4" gridrow="0" pageId="80" pageNumber="81">Exemplar transformation series</th>
|
||
<th box="[1326,1530,437,517]" gridcol="5" gridrow="0" pageId="80" pageNumber="81">
|
||
<emphasis bold="true" box="[1326,1417,437,458]" pageId="80" pageNumber="81">Excluded</emphasis>
|
||
<emphasis bold="true" box="[1326,1478,466,517]" pageId="80" pageNumber="81">transformation series</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[108,1530,536,588]" gridrow="1" pageId="80" pageNumber="81">
|
||
<th box="[108,184,536,588]" gridcol="0" gridrow="1" pageId="80" pageNumber="81">5</th>
|
||
<td box="[227,380,536,588]" gridcol="1" gridrow="1" pageId="80" pageNumber="81">Skull height</td>
|
||
<td box="[422,709,536,588]" gridcol="2" gridrow="1" pageId="80" pageNumber="81">4, 57, 66, 238, 239, 240, 254, 260, 570</td>
|
||
<td box="[744,1032,536,588]" gridcol="3" gridrow="1" pageId="80" pageNumber="81">4, 66; 570; 57, 238, 239, 240; 260</td>
|
||
<td box="[1067,1282,536,588]" gridcol="4" gridrow="1" pageId="80" pageNumber="81">4, 240, 260, 570</td>
|
||
<td box="[1326,1530,536,588]" gridcol="5" gridrow="1" pageId="80" pageNumber="81">66; 57, 238, 239</td>
|
||
</tr>
|
||
<tr box="[108,1530,603,625]" gridrow="2" pageId="80" pageNumber="81">
|
||
<th box="[108,184,603,625]" gridcol="0" gridrow="2" pageId="80" pageNumber="81">5</th>
|
||
<td box="[227,380,603,625]" gridcol="1" gridrow="2" pageId="80" pageNumber="81">Antorbital fossa</td>
|
||
<td box="[422,709,603,625]" gridcol="2" gridrow="2" pageId="80" pageNumber="81">374, 375</td>
|
||
<td box="[744,1032,603,625]" gridcol="3" gridrow="2" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1067,1282,603,625]" gridcol="4" gridrow="2" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1326,1530,603,625]" gridcol="5" gridrow="2" pageId="80" pageNumber="81">n/a</td>
|
||
</tr>
|
||
<tr box="[108,1530,641,692]" gridrow="3" pageId="80" pageNumber="81">
|
||
<th box="[108,184,641,692]" gridcol="0" gridrow="3" pageId="80" pageNumber="81">5</th>
|
||
<td box="[227,380,641,692]" gridcol="1" gridrow="3" pageId="80" pageNumber="81">Dorsal margin of postorbital</td>
|
||
<td box="[422,709,641,692]" gridcol="2" gridrow="3" pageId="80" pageNumber="81">620, 623</td>
|
||
<td box="[744,1032,641,692]" gridcol="3" gridrow="3" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1067,1282,641,692]" gridcol="4" gridrow="3" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1326,1530,641,692]" gridcol="5" gridrow="3" pageId="80" pageNumber="81">n/a</td>
|
||
</tr>
|
||
<tr box="[108,1530,708,759]" gridrow="4" pageId="80" pageNumber="81">
|
||
<th box="[108,184,708,759]" gridcol="0" gridrow="4" pageId="80" pageNumber="81">5</th>
|
||
<td box="[227,380,708,759]" gridcol="1" gridrow="4" pageId="80" pageNumber="81">Subcutaneous surface</td>
|
||
<td box="[422,709,708,759]" gridcol="2" gridrow="4" pageId="80" pageNumber="81">556, 621</td>
|
||
<td box="[744,1032,708,759]" gridcol="3" gridrow="4" pageId="80" pageNumber="81">556, 621</td>
|
||
<td box="[1067,1282,708,759]" gridcol="4" gridrow="4" pageId="80" pageNumber="81">621</td>
|
||
<td box="[1326,1530,708,759]" gridcol="5" gridrow="4" pageId="80" pageNumber="81">556</td>
|
||
</tr>
|
||
<tr box="[108,1530,774,825]" gridrow="5" pageId="80" pageNumber="81">
|
||
<th box="[108,184,774,825]" gridcol="0" gridrow="5" pageId="80" pageNumber="81">5</th>
|
||
<td box="[227,380,774,825]" gridcol="1" gridrow="5" pageId="80" pageNumber="81">Maxillary tooth count</td>
|
||
<td box="[422,709,774,825]" gridcol="2" gridrow="5" pageId="80" pageNumber="81">132, 397, 1176</td>
|
||
<td box="[744,1032,774,825]" gridcol="3" gridrow="5" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1067,1282,774,825]" gridcol="4" gridrow="5" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1326,1530,774,825]" gridcol="5" gridrow="5" pageId="80" pageNumber="81">n/a</td>
|
||
</tr>
|
||
<tr box="[108,1530,841,951]" gridrow="6" pageId="80" pageNumber="81">
|
||
<th box="[108,184,841,951]" gridcol="0" gridrow="6" pageId="80" pageNumber="81">6</th>
|
||
<td box="[227,380,841,951]" gridcol="1" gridrow="6" pageId="80" pageNumber="81">Lacrimal inflation</td>
|
||
<td box="[422,709,841,951]" gridcol="2" gridrow="6" pageId="80" pageNumber="81">38, 40, 41, 45, 46, 436, 437, 447, 450, 453, 454, 459, 461, 462, 464, 465, 468, 471, 482, 485</td>
|
||
<td box="[744,1032,841,951]" gridcol="3" gridrow="6" pageId="80" pageNumber="81">40, 45, 436, 447, 454, 462, 465, 468; 46; 437, 450; 464; 453; 471; 459; 41, 461; 39; 38, 482, 485</td>
|
||
<td box="[1067,1282,841,951]" gridcol="4" gridrow="6" pageId="80" pageNumber="81">447; 46; 450; 464; 453; 471; 459; 41; 39; 38</td>
|
||
<td box="[1326,1530,841,951]" gridcol="5" gridrow="6" pageId="80" pageNumber="81">40, 45, 436, 454, 462, 465, 468; 437; 461; 482, 485</td>
|
||
</tr>
|
||
<tr box="[108,1530,966,988]" gridrow="7" pageId="80" pageNumber="81">
|
||
<th box="[108,184,966,988]" gridcol="0" gridrow="7" pageId="80" pageNumber="81">6</th>
|
||
<td box="[227,380,966,988]" gridcol="1" gridrow="7" pageId="80" pageNumber="81">Skull frame</td>
|
||
<td box="[422,709,966,988]" gridcol="2" gridrow="7" pageId="80" pageNumber="81">517, 522, 626</td>
|
||
<td box="[744,1032,966,988]" gridcol="3" gridrow="7" pageId="80" pageNumber="81">517, 626; 522</td>
|
||
<td box="[1067,1282,966,988]" gridcol="4" gridrow="7" pageId="80" pageNumber="81">517; 522</td>
|
||
<td box="[1326,1530,966,988]" gridcol="5" gridrow="7" pageId="80" pageNumber="81">626</td>
|
||
</tr>
|
||
<tr box="[108,1530,1004,1055]" gridrow="8" pageId="80" pageNumber="81">
|
||
<th box="[108,184,1004,1055]" gridcol="0" gridrow="8" pageId="80" pageNumber="81">7</th>
|
||
<td box="[227,380,1004,1055]" gridcol="1" gridrow="8" pageId="80" pageNumber="81">Lacrimal inflation</td>
|
||
<td box="[422,709,1004,1055]" gridcol="2" gridrow="8" pageId="80" pageNumber="81">39, 41, 43, 453</td>
|
||
<td box="[744,1032,1004,1055]" gridcol="3" gridrow="8" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1067,1282,1004,1055]" gridcol="4" gridrow="8" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1326,1530,1004,1055]" gridcol="5" gridrow="8" pageId="80" pageNumber="81">n/a</td>
|
||
</tr>
|
||
<tr box="[108,1530,1071,1093]" gridrow="9" pageId="80" pageNumber="81">
|
||
<th box="[108,184,1071,1093]" gridcol="0" gridrow="9" pageId="80" pageNumber="81">9</th>
|
||
<td box="[227,380,1071,1093]" gridcol="1" gridrow="9" pageId="80" pageNumber="81">Alveolus size</td>
|
||
<td box="[422,709,1071,1093]" gridcol="2" gridrow="9" pageId="80" pageNumber="81">117, 121</td>
|
||
<td box="[744,1032,1071,1093]" gridcol="3" gridrow="9" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1067,1282,1071,1093]" gridcol="4" gridrow="9" pageId="80" pageNumber="81">n/a</td>
|
||
<td box="[1326,1530,1071,1093]" gridcol="5" gridrow="9" pageId="80" pageNumber="81">n/a</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="80.[498,1542,1168,1712]" pageId="80" pageNumber="81">
|
||
The analysis recovered 20,000 MPTs (the limit of memory capacity); a strict consensus ontogram shows several polytomies among juveniles (
|
||
<materialsCitation box="[1259,1497,1447,1474]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47787" pageId="80" pageNumber="81" specimenCode="AMNH FARB 5050">AMNH FARB 5050</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,652,1487,1514]" collectionCode="CMNH" pageId="80" pageNumber="81" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
,
|
||
<materialsCitation box="[663,802,1487,1514]" collectionCode="DDM" pageId="80" pageNumber="81" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[813,974,1487,1513]" collectionCode="LACM" pageId="80" pageNumber="81" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
), subadults (BMRP 2006.6.4,
|
||
<materialsCitation box="[1314,1473,1487,1514]" collectionCode="LACM" pageId="80" pageNumber="81" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="RSM" pageId="80" pageNumber="81" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
), and adults distal to
|
||
<materialsCitation box="[834,967,1527,1554]" collectionCode="MOR" pageId="80" pageNumber="81" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
. Within the adult polytomy, one sister pair was recovered (
|
||
<materialsCitation box="[626,742,1567,1594]" collectionCode="MOR" pageId="80" pageNumber="81" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
+
|
||
<materialsCitation box="[772,892,1567,1593]" collectionCode="MOR" pageId="80" pageNumber="81" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
). These regions of the ontogram have low Bremer (1
|
||
<emphasis box="[1496,1511,1568,1593]" italics="true" pageId="80" pageNumber="81">–</emphasis>
|
||
4), but relatively high bootstrap (62
|
||
<emphasis box="[878,893,1607,1632]" italics="true" pageId="80" pageNumber="81">–</emphasis>
|
||
97), and jackknife (61
|
||
<emphasis box="[1150,1165,1607,1632]" italics="true" pageId="80" pageNumber="81">–</emphasis>
|
||
97) values, indicating that the conflicting results are almost certainly an artifact character removal that decreased the proportion of topologically informative data.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection box="[498,728,1753,1788]" pageId="80" pageNumber="81" type="discussion">
|
||
<paragraph blockId="80.[498,728,1753,1788]" box="[498,728,1753,1788]" pageId="80" pageNumber="81">
|
||
<heading allCaps="true" bold="true" box="[498,728,1753,1788]" fontSize="14" level="1" pageId="80" pageNumber="81" reason="0">
|
||
<emphasis bold="true" box="[498,728,1753,1788]" pageId="80" pageNumber="81">DISCUSSION</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection box="[498,980,1808,1838]" pageId="80" pageNumber="81" type="nomenclature">
|
||
<paragraph blockId="80.[498,1542,1808,1920]" box="[498,980,1808,1838]" pageId="80" pageNumber="81">
|
||
<heading bold="true" box="[498,980,1808,1838]" fontSize="12" level="2" pageId="80" pageNumber="81" reason="0">
|
||
<emphasis bold="true" box="[498,980,1808,1838]" pageId="80" pageNumber="81">
|
||
Diagnosis of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[698,980,1808,1838]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="80" pageNumber="81" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[698,980,1808,1838]" italics="true" pageId="80" pageNumber="81">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="93" lastPageNumber="94" pageId="80" pageNumber="81" type="discussion">
|
||
<paragraph blockId="80.[498,1542,1808,1920]" lastBlockId="81.[498,1542,234,1576]" lastPageId="81" lastPageNumber="82" pageId="80" pageNumber="81">
|
||
The ontogenetic position of
|
||
<materialsCitation box="[833,943,1854,1880]" collectionCode="CM" pageId="80" pageNumber="81" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
among the most mature specimens in the sample (
|
||
<figureCitation box="[508,582,1893,1920]" captionStart="Figure 1" captionStartId="9.[116,181,1249,1271]" captionTargetBox="[90,1542,234,1236]" captionTargetId="graphics-216@9.[341,1528,267,1221]" captionTargetPageId="9" captionText="Figure 1 Results of the cladistic analysis of 1,850 characters among 44 specimens of Tyrannosaurus rex. (A) Strict consensus of 50 MPTs showing the recovery of three primary growth stages separated by the specimen BMRP 2002.4.1. (B) The single ontogram recovered after the exclusion of wildcard specimens, reducing the number of OTUs to 31. Numbers to the left of the internodes are bootstrap and jackknife values, respectively; numbers to the right are Bremer decay indices. Asterisk indicates the type specimen. Ellipses enclose the regions of polytomies produced by the wildcard specimens, which are listed in the lower right hand corner of the corresponding ellipse. Note that the ellipses are limited to one side or the other relative to BMRP 2002.4.1, which corresponds to the topology of the strict consensus ontogram. Full-size DOI: 10.7717/peerj.9192/fig-1" figureDoi="http://doi.org/10.5281/zenodo.5824974" httpUri="https://zenodo.org/record/5824974/files/figure.png" pageId="80" pageNumber="81">Figs. 1</figureCitation>
|
||
and
|
||
<figureCitation box="[639,653,1894,1920]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="80" pageNumber="81">2</figureCitation>
|
||
) indicates that the taxon,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[953,1020,1894,1919]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="80" pageNumber="81" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[953,1020,1894,1919]" italics="true" pageId="80" pageNumber="81">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, is indeed based upon a mature adult and, by extension, the type is a defensible name bearer, foundation for diagnosis, and point of comparison for other specimens. The results here support one of the diagnostic features of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[530,599,315,340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[530,599,315,340]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
first identified by
|
||
<bibRefCitation author="Osborn HF" box="[818,984,314,340]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="81" pageNumber="82" pagination="259 - 265" part="21" refId="ref55417" refString="Osborn HF. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 21: 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">
|
||
<emphasis box="[818,984,314,340]" italics="true" pageId="81" pageNumber="82">Osborn (1905)</emphasis>
|
||
</bibRefCitation>
|
||
, namely gigantic size. The other characters are not diagnostic;
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[722,790,355,380]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[722,790,355,380]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
did not have a long humerus and the absence of armor plates is symplesiomorphic for
|
||
<taxonomicName box="[764,964,394,420]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
, if not Ornthodira.
|
||
<bibRefCitation author="Osborn HF" box="[1199,1365,394,420]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="81" pageNumber="82" pagination="259 - 265" part="21" refId="ref55417" refString="Osborn HF. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 21: 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">
|
||
<emphasis box="[1199,1365,394,420]" italics="true" pageId="81" pageNumber="82">Osborn (1905)</emphasis>
|
||
</bibRefCitation>
|
||
observed that
|
||
<emphasis box="[553,851,434,459]" italics="true" pageId="81" pageNumber="82">Dynamosaurus imperious</emphasis>
|
||
(junior subjective synonym of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1220,1289,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1220,1289,434,459]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
) was distinguished by its number of dentary teeth
|
||
<emphasis box="[856,867,473,499]" italics="true" pageId="81" pageNumber="82">“</emphasis>
|
||
twelve to thirteen
|
||
<emphasis box="[1070,1081,473,499]" italics="true" pageId="81" pageNumber="82">”</emphasis>
|
||
; indeed, the results here find that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1475,1542,474,499]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1475,1542,474,499]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
has the lowest number of dentary teeth (12) among tyrannosaurids. The same is also true of the maxilla, which has as few as 11 teeth (e.g.,
|
||
<materialsCitation box="[1083,1247,553,579]" collectionCode="LACM" pageId="81" pageNumber="82" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="81.[498,1542,234,1576]" pageId="81" pageNumber="82">
|
||
In 1906, Osborn synonymized
|
||
<taxonomicName box="[895,1054,594,619]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="imperiosus">
|
||
<emphasis box="[895,1054,594,619]" italics="true" pageId="81" pageNumber="82">D. imperiosus</emphasis>
|
||
</taxonomicName>
|
||
with
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1124,1193,594,619]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1124,1193,594,619]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and he expanded the list of diagnostic characters. Of these, several are symplesiomorphies: presence of the internal antorbital fenestra and the maxillary fenestra, horizontal bar of the squamosal, presence of the subnarial foramen, small first dentary tooth, presence of interdental plates, 23 presacral vertebrae, five sacral vertebrae, sacral spinous processes fused into a single plate, atlantoaxial complex composed of six segments, reduced scapula and humerus, presence of abdominal ribs, pelvic construction, hollow limb and girdle bones, and long hind limbs.
|
||
</paragraph>
|
||
<paragraph blockId="81.[498,1542,234,1576]" pageId="81" pageNumber="82">
|
||
Some characters (
|
||
<bibRefCitation author="Osborn HF" box="[740,895,872,898]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="81" pageNumber="82" pagination="281 - 296" part="22" refId="ref55443" refString="Osborn HF. 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication). Bulletin of the American Museum of Natural History 22: 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication)" type="journal article" year="1906">
|
||
<emphasis box="[740,895,872,898]" italics="true" pageId="81" pageNumber="82">Osborn, 1906</emphasis>
|
||
</bibRefCitation>
|
||
) are mischaracterizations, such as the abbreviated skull (whereas it is dorsoventrally deep, not rostrocaudally short) and the primary metatarsals partly co-ossified (although tightly apposed, these bones are separate structures). Also, his observation of teeth in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1022,1091,992,1017]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1022,1091,992,1017]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
that are
|
||
<emphasis box="[1198,1209,991,1017]" italics="true" pageId="81" pageNumber="82">“</emphasis>
|
||
very broadly oval in section, transverse exceeding anteroposterior diameters
|
||
<emphasis box="[1050,1062,1031,1057]" italics="true" pageId="81" pageNumber="82">”</emphasis>
|
||
(
|
||
<bibRefCitation author="Osborn HF" box="[1080,1296,1031,1058]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="81" pageNumber="82" pagination="281 - 296" part="22" refId="ref55443" refString="Osborn HF. 1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication). Bulletin of the American Museum of Natural History 22: 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur: (second communication)" type="journal article" year="1906">
|
||
<emphasis box="[1080,1235,1031,1057]" italics="true" pageId="81" pageNumber="82">Osborn, 1906</emphasis>
|
||
: 283
|
||
</bibRefCitation>
|
||
), deserves comment. A comparison of width to length ratios among adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1126,1195,1072,1097]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1126,1195,1072,1097]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and representative adults of
|
||
<taxonomicName box="[528,757,1111,1137]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis box="[528,703,1111,1137]" italics="true" pageId="81" pageNumber="82">Daspletosaurus</emphasis>
|
||
spp.
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[813,986,1111,1137]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis box="[813,986,1111,1137]" italics="true" pageId="81" pageNumber="82">A. sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
(aside from the first two maxillary teeth and the first dentary tooth, which tend to be incisiform or conical and so incomparable with so-called lateral teeth that are ziphiform in shape) shows that dentary teeth that are wider than long have a scattered distribution among
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1051,1120,1231,1256]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1051,1120,1231,1256]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
adults (
|
||
<tableCitation box="[1217,1319,1230,1257]" captionStart="Table 11" captionStartId="48.[117,172,248,269]" captionTargetBox="[108,1525,349,1723]" captionText="Table 11 Summary of width to length ratios of maxillary and dentary teeth in adult Tyrannosaurus rex compared with representative adults of other tyrannosaurids. In T. rex the maxillary teeth tend to be wider than in other tyrannosaurids, and also at the mesial end of the tooth row of the dentary. However, the characterization by Osborn (1906) of the teeth in T. rex as generally wider than long is not supported by these data." httpUri="http://table.plazi.org/id/DF5C84E2FFF4FF98FF93FF7032AAFEF5" pageId="81" pageNumber="82" tableUuid="DF5C84E2FFF4FF98FF93FF7032AAFEF5">Table 11</tableCitation>
|
||
). Maxillary teeth with widths that exceed their lengths are not seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1126,1195,1271,1296]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1126,1195,1271,1296]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, but, in general, the maxillary teeth of adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[782,851,1311,1336]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[782,851,1311,1336]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
tend to be wider than what is seen in its close relatives, and the dentary teeth are somewhat narrower (
|
||
<tableCitation box="[1057,1158,1350,1376]" captionStart="Table 11" captionStartId="48.[117,172,248,269]" captionTargetBox="[108,1525,349,1723]" captionText="Table 11 Summary of width to length ratios of maxillary and dentary teeth in adult Tyrannosaurus rex compared with representative adults of other tyrannosaurids. In T. rex the maxillary teeth tend to be wider than in other tyrannosaurids, and also at the mesial end of the tooth row of the dentary. However, the characterization by Osborn (1906) of the teeth in T. rex as generally wider than long is not supported by these data." httpUri="http://table.plazi.org/id/DF5C84E2FFF4FF98FF93FF7032AAFEF5" pageId="81" pageNumber="82" tableUuid="DF5C84E2FFF4FF98FF93FF7032AAFEF5">Table 11</tableCitation>
|
||
). In contrast, the maximum width to length ratio of lateral teeth (excluding wide mesial teeth) in young adults, such as
|
||
<materialsCitation box="[593,725,1429,1456]" collectionCode="MOR" pageId="81" pageNumber="82" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, is only 68% (mean of all
|
||
<materialsCitation box="[1035,1165,1429,1456]" collectionCode="MOR" pageId="81" pageNumber="82" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
teeth: 72%), indicating that autapomorphically wide teeth occur late in growth. Therefore, only three of Osborn
|
||
<emphasis box="[1487,1493,1470,1496]" italics="true" pageId="81" pageNumber="82">’</emphasis>
|
||
s diagnostic characters are seen here: gigantic size, the low count of twelve dentary teeth, and the extreme width of the maxillary- and mesial dentary teeth.
|
||
</paragraph>
|
||
<paragraph blockId="81.[498,1542,1596,1865]" box="[498,760,1596,1624]" pageId="81" pageNumber="82">
|
||
<heading bold="true" box="[498,760,1596,1624]" fontSize="11" level="3" pageId="81" pageNumber="82" reason="6">
|
||
<emphasis bold="true" box="[498,760,1596,1624]" italics="true" pageId="81" pageNumber="82">Individual variation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="81.[498,1542,1596,1865]" lastBlockId="82.[498,1542,234,978]" lastPageId="82" lastPageNumber="83" pageId="81" pageNumber="82">
|
||
In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[533,601,1640,1665]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="81" pageNumber="82" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[533,601,1640,1665]" italics="true" pageId="81" pageNumber="82">T. rex</emphasis>
|
||
</taxonomicName>
|
||
individual variation increases with maturity, which contradicts the naïve expectation that variation should decrease upon the passage through an accumulation of developmental constraints throughout growth (see
|
||
<emphasis box="[1089,1100,1719,1745]" italics="true" pageId="81" pageNumber="82">“</emphasis>
|
||
Results
|
||
<emphasis box="[1183,1195,1719,1745]" italics="true" pageId="81" pageNumber="82">”</emphasis>
|
||
above). Evidently, constraints are tightest early in growth, as shown by the absence of character reversals from juveniles and the least mature young adult. The presence of reversals throughout adulthood is evidence that variation is less tightly constrained, allowing characters to freely reverse and progress. Also, the onset of reversals coincides with crossing the plesiomorphic 3,000 kg threshold in mass (
|
||
<figureCitation box="[834,899,274,300]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="82" pageNumber="83">Fig. 2</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="82.[498,1542,234,978]" pageId="82" pageNumber="83">
|
||
The presence of more reversals than progressions in
|
||
<materialsCitation box="[1154,1340,314,340]" collectionCode="FMNH" pageId="82" pageNumber="83" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
, presumably the most mature specimen in the sample, raises the question of whether or not an individual specimen should be considered the heuristic representative of the terminal growth stage on the ontogram. Among adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1024,1092,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1024,1092,434,459]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, each node is supported by a relatively low number of synontomorphies that include progressions and reversals; the characters that diagnose
|
||
<materialsCitation box="[664,848,513,540]" collectionCode="FMNH" pageId="82" pageNumber="83" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
are dominated by reversals, which is the pattern seen in the individual variation of other adults (
|
||
<materialsCitation box="[972,1209,553,580]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="82" pageNumber="83" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1222,1458,553,580]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47763" pageId="82" pageNumber="83" specimenCode="AMNH FARB 5029">AMNH FARB 5029</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,734,593,620]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="82" pageNumber="83" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
,
|
||
<materialsCitation box="[747,908,593,619]" collectionCode="LACM" pageId="82" pageNumber="83" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<materialsCitation box="[921,1039,593,620]" collectionCode="MOR" pageId="82" pageNumber="83" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1053,1171,593,619]" collectionCode="MOR" pageId="82" pageNumber="83" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1184,1316,593,619]" collectionCode="MOR" pageId="82" pageNumber="83" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1329,1461,593,619]" collectionCode="MOR" pageId="82" pageNumber="83" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
,
|
||
<materialsCitation box="[498,709,633,659]" collectionCode="NMMNH" pageId="82" pageNumber="83" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
,
|
||
<materialsCitation box="[722,867,632,659]" collectionCode="RSM" pageId="82" pageNumber="83" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
,
|
||
<materialsCitation box="[880,1035,633,659]" collectionCode="SDSM" pageId="82" pageNumber="83" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1048,1221,633,659]" collectionCode="UWBM" pageId="82" pageNumber="83" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
, UWGM 191). Among synontomorphies, where both reversals and progressions are seen, in only one case do the progressions outnumber the reversals, which emphasizes the difference seen in
|
||
<materialsCitation collectionCode="FMNH" pageId="82" pageNumber="83" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="82.[498,1542,234,978]" pageId="82" pageNumber="83">It is possible that the reversals might have been actual reversals over the lifetime of the individual (e.g., 0 to 1 to 0), or juvenile characters that were carried unchanged into adulthood. However, it is not possible to make the distinction between reversals and static character states of individuals, on the one hand, from true ontogenetic reversals on the other, without knowing the entire ontogeny of the individual organism in question.</paragraph>
|
||
<paragraph blockId="82.[498,1542,1017,1884]" box="[498,713,1017,1045]" pageId="82" pageNumber="83">
|
||
<heading bold="true" box="[498,713,1017,1045]" fontSize="11" level="3" pageId="82" pageNumber="83" reason="6">
|
||
<emphasis bold="true" box="[498,713,1017,1045]" italics="true" pageId="82" pageNumber="83">Sexual maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="82.[498,1542,1017,1884]" pageId="82" pageNumber="83">
|
||
Among extant archosaurs (e.g.,
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[873,1159,1060,1086]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[873,1159,1060,1086]" italics="true" pageId="82" pageNumber="83">Alligator mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
), sexual maturity is reached at approximately half adult size (
|
||
<bibRefCitation author="Wilkinson PM & Rhodes WE" box="[847,1150,1100,1127]" journalOrPublisher="Journal of Wildlife Management" pageId="82" pageNumber="83" pagination="397 - 402" part="61" refId="ref56571" refString="Wilkinson PM, Rhodes WE. 1997. Growth rates of American alligators in coastal South Carolina. Journal of Wildlife Management 61 (2): 397 - 402 DOI 10.2307 / 3802596." title="Growth rates of American alligators in coastal South Carolina" type="journal article" year="1997">
|
||
<emphasis box="[847,1150,1100,1127]" italics="true" pageId="82" pageNumber="83">Wilkinson & Rhodes, 1997</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Fernandez Blanco MV & Bona P & Scheyer TM" box="[1162,1343,1100,1126]" journalOrPublisher="Journal of Morphology" pageId="82" pageNumber="83" pagination="259 - 273" part="279" refId="ref53510" refString="Foth C, Fernandez Blanco MV, Bona P, Scheyer TM. 2018. Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri. Journal of Morphology 279 (2): 259 - 273 DOI 10.1002 / jmor. 20769." title="Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri" type="journal article" year="2018">
|
||
<emphasis box="[1162,1343,1100,1126]" italics="true" pageId="82" pageNumber="83">Foth et al., 2018</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Wilkinson PM & Rainwater TR & Woodward AR & Leone EH & Carter C." journalOrPublisher="Copeia" pageId="82" pageNumber="83" pagination="843 - 852" part="104" refId="ref56518" refString="Wilkinson PM, Rainwater TR, Woodward AR, Leone EH, Carter C. 2016. Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures. Copeia 104 (4): 843 - 852 DOI 10.1643 / CH- 16 - 430." title="Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures" type="journal article" year="2016">
|
||
<emphasis italics="true" pageId="82" pageNumber="83">Wilkinson et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
). If this pattern is plesiomorphic for Archosauria (
|
||
<taxonomicName box="[1154,1284,1140,1166]" class="Reptilia" kingdom="Animalia" order="Crocodylia" pageId="82" pageNumber="83" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
|
||
+
|
||
<taxonomicName authorityName="Owen" authorityYear="1842" box="[1318,1451,1140,1167]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
), the naïve null hypothesis for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[799,867,1181,1206]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[799,867,1181,1206]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is that the onset of sexual maturity occurred when skull length reached 70 cm. Ergo, specimens such as
|
||
<materialsCitation box="[1061,1244,1220,1246]" collectionCode="BMRP" pageId="82" pageNumber="83" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
(skull length: 74 cm) indicate that sexual maturity occurred before the onset of the extreme transition from long and low skulls to the deep and stocky frame of subadults and adults (
|
||
<figureCitation box="[1380,1460,1299,1326]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="82" pageNumber="83">Fig. 12</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="82.[498,1542,1017,1884]" pageId="82" pageNumber="83">
|
||
At the very least, it is improbable that the 15-year-old female subadult
|
||
<materialsCitation box="[1360,1542,1339,1366]" collectionCode="BMRP" pageId="82" pageNumber="83" specimenCode="BMRP 2006.4.4">BMRP 2006.4.4</materialsCitation>
|
||
represents the earliest onset of sexual maturity in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1076,1432,1379,1406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1076,1143,1380,1405]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
(
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[1160,1421,1379,1405]" journalOrPublisher="Science Advances" pageId="82" pageNumber="83" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[1160,1421,1379,1405]" italics="true" pageId="82" pageNumber="83">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. If sexual maturity occurred earlier, then
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[869,938,1420,1445]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[869,938,1420,1445]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is an organism with determinate growth in which growth continues after the onset of sexual maturation and stops before senescence (sensu
|
||
<bibRefCitation author="Lincoln R & Boxshall G & Clark P." box="[579,942,1499,1525]" journalOrPublisher="London: Cambridge University Press" pageId="82" pageNumber="83" refId="ref54943" refString="Lincoln R, Boxshall G, Clark P. 1982. A dictionary of ecology, evolution, and sytematics. Second Edition. London: Cambridge University Press." title="A dictionary of ecology, evolution, and sytematics. Second Edition" type="book" year="1982">
|
||
<emphasis box="[579,942,1499,1525]" italics="true" pageId="82" pageNumber="83">Lincoln, Boxshall & Clark, 1982</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Wilkinson PM & Rainwater TR & Woodward AR & Leone EH & Carter C." box="[978,1238,1499,1525]" journalOrPublisher="Copeia" pageId="82" pageNumber="83" pagination="843 - 852" part="104" refId="ref56518" refString="Wilkinson PM, Rainwater TR, Woodward AR, Leone EH, Carter C. 2016. Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures. Copeia 104 (4): 843 - 852 DOI 10.1643 / CH- 16 - 430." title="Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures" type="journal article" year="2016">
|
||
<emphasis box="[978,1238,1499,1525]" italics="true" pageId="82" pageNumber="83">Wilkinson et al. (2016)</emphasis>
|
||
</bibRefCitation>
|
||
). As in
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[1327,1536,1499,1525]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[1327,1536,1499,1525]" italics="true" pageId="82" pageNumber="83">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
, the growth rate in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[720,789,1539,1564]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[720,789,1539,1564]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
reduces once sexual maturity is reached, where it is relatively low in large juveniles and subadults (
|
||
<bibRefCitation author="Woodward H & Tremaine K & Williams SA & Zanno LE & Horner JR & Myhrvold N." box="[938,1203,1578,1605]" journalOrPublisher="Science Advances" pageId="82" pageNumber="83" pagination="eaax 6250" part="6" refId="ref56786" refString="Woodward H, Tremaine K, Williams SA, Zanno LE, Horner JR, Myhrvold N. 2020. Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Science Advances 6 (1): eaax 6250 DOI 10.1126 / sciadv. aax 6250." title="Growing up Tyrannosaurus rex: osteohistology refutes the pygmy Nanotyrannus and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">
|
||
<emphasis box="[938,1203,1578,1605]" italics="true" pageId="82" pageNumber="83">Woodward et al., 2020</emphasis>
|
||
</bibRefCitation>
|
||
) and it nearly ceases in the young adults and adult categories (
|
||
<bibRefCitation author="Horner JR & Padian K." box="[913,1189,1618,1645]" journalOrPublisher="Proceedings of the Royal Society of London. Series B: Biological Sciences" pageId="82" pageNumber="83" pagination="1875 - 1880" part="271" refId="ref54442" refString="Horner JR, Padian K. 2004. Age and growth dynamics of Tyrannosaurus rex. Proceedings of the Royal Society of London. Series B: Biological Sciences 271 (1551): 1875 - 1880 DOI 10.1098 / rspb. 2004.2829." title="Age and growth dynamics of Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[913,1189,1618,1645]" italics="true" pageId="82" pageNumber="83">Horner & Padian, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). If these patterns of growth rate are comparable, then it is plesiomorphic and unaffected by metabolism (i.e., ectothermy in crocodylians, endothermy in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1063,1130,1699,1724]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1063,1130,1699,1724]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
). It is thought that the pattern seen in
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[529,737,1738,1764]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[529,737,1738,1764]" italics="true" pageId="82" pageNumber="83">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
evolved as an energy expense-optimization strategy (
|
||
<bibRefCitation author="Wilkinson PM & Rainwater TR & Woodward AR & Leone EH & Carter C." journalOrPublisher="Copeia" pageId="82" pageNumber="83" pagination="843 - 852" part="104" refId="ref56518" refString="Wilkinson PM, Rainwater TR, Woodward AR, Leone EH, Carter C. 2016. Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures. Copeia 104 (4): 843 - 852 DOI 10.1643 / CH- 16 - 430." title="Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures" type="journal article" year="2016">
|
||
<emphasis italics="true" pageId="82" pageNumber="83">Wilkinson et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
), which might be the case for all nonavian archosaurs. Given that female
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[498,709,1818,1844]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[498,709,1818,1844]" italics="true" pageId="82" pageNumber="83">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
can produce viable eggs for their entire lives, the term
|
||
<emphasis box="[1363,1374,1818,1844]" italics="true" pageId="82" pageNumber="83">“</emphasis>
|
||
senescent
|
||
<emphasis box="[1485,1497,1818,1844]" italics="true" pageId="82" pageNumber="83">”</emphasis>
|
||
as used here for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[662,731,1858,1883]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="82" pageNumber="83" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[662,731,1858,1883]" italics="true" pageId="82" pageNumber="83">T. rex</emphasis>
|
||
</taxonomicName>
|
||
does not imply that senescent adults are nonreproductive.
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,233,1419]" box="[498,765,233,261]" pageId="83" pageNumber="84">
|
||
<heading bold="true" box="[498,765,233,261]" fontSize="11" level="3" pageId="83" pageNumber="84" reason="6">
|
||
<emphasis bold="true" box="[498,765,233,261]" italics="true" pageId="83" pageNumber="84">Sexual dimorphism</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,233,1419]" pageId="83" pageNumber="84">
|
||
Sexual dimorphism is marked by one or more distinct phenotypic differences between males and females, excluding differences in size (
|
||
<bibRefCitation author="Padian K & Horner JR" box="[1061,1331,316,343]" journalOrPublisher="Journal of Zoology" pageId="83" pageNumber="84" pagination="23 - 27" part="283" refId="ref55513" refString="Padian K, Horner JR. 2011. The definition of sexual selection and its implications for dinosaurian biology. Journal of Zoology 283 (1): 23 - 27 DOI 10.1111 / j. 1469 - 7998.2010.00761. x." title="The definition of sexual selection and its implications for dinosaurian biology" type="journal article" year="2011">
|
||
<emphasis box="[1061,1331,316,343]" italics="true" pageId="83" pageNumber="84">Padian & Horner, 2011</emphasis>
|
||
</bibRefCitation>
|
||
). The inference of the absence of sexual dimorphism in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[940,1009,357,382]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[940,1009,357,382]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is supported by three lines of evidence: (1) the ontogram is linear and pectinate, lacking a distinct bifurcation that unites females on one branch and males on the other; (2) the number of specimens more mature than
|
||
<materialsCitation box="[498,629,476,503]" collectionCode="MOR" pageId="83" pageNumber="84" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
is high (22), indicating that (a) the sample size is sufficient to capture the signal of two distinct morphs and (b) the probability of sampling only one sex over the other is vanishingly low; and (3) individual variation does not separate out two identifiable groups among subadult and adult specimens. In other words, aside from subtle differences, all adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[743,812,636,661]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[743,812,636,661]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
simply look alike.
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,233,1419]" pageId="83" pageNumber="84">
|
||
The absence of dimorphism pertains to the osteodental characters included in the analysis; in life, dimorphism might have been expressed by differences in soft tissues (e.g., integument pigmentation) that do not preserve as fossils. The possibility that dimorphism was expressed by size and growth rate, as is seen in living crocodylians (
|
||
<bibRefCitation author="Taylor P & Li F & Holland A & Martin M & Rosenblatt AE" box="[508,721,834,861]" journalOrPublisher="Amphibia-Reptilia" pageId="83" pageNumber="84" pagination="9 - 14" part="27" refId="ref56224" refString="Taylor P, Li F, Holland A, Martin M, Rosenblatt AE. 2016. Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana. Amphibia-Reptilia 27 (1): 9 - 14 DOI 10.1163 / 15685381 - 00003024." title="Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana" type="journal article" year="2016">
|
||
<emphasis box="[508,721,834,861]" italics="true" pageId="83" pageNumber="84">Taylor et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
), requires a sample of unambiguously female specimens, which is currently unavailable for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[796,865,875,900]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[796,865,875,900]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Finally, it is notable that among living crocodylians (e.g.,
|
||
<taxonomicName authorityName="Daudin" authorityYear="1802" box="[562,670,915,941]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="yacare">
|
||
<emphasis box="[562,670,915,941]" italics="true" pageId="83" pageNumber="84">C. yacare</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Cogger" authorityYear="2000" box="[685,928,914,940]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="johnstoni">
|
||
<emphasis box="[685,928,914,940]" italics="true" pageId="83" pageNumber="84">Crocodylus johnstoni</emphasis>
|
||
</taxonomicName>
|
||
) dimorphism in size is seen between hatchlings, where females are smaller than males (
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[958,1187,954,981]" journalOrPublisher="PLOS ONE" pageId="83" pageNumber="84" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[958,1187,954,981]" italics="true" pageId="83" pageNumber="84">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Edwards GP & Webb GJ & Manolis SC & Mazanov A." box="[1202,1438,954,980]" journalOrPublisher="Australian Journal of Zoology" pageId="83" pageNumber="84" pagination="97" part="65" refId="ref53078" refString="Edwards GP, Webb GJ, Manolis SC, Mazanov A. 2017. Morphometric analysis of the Australian freshwater crocodile (Crocodylus johnstoni). Australian Journal of Zoology 65 (2): 97 DOI 10.1071 / ZO 16079." title="Morphometric analysis of the Australian freshwater crocodile (Crocodylus johnstoni)" type="journal article" year="2017">
|
||
<emphasis box="[1202,1438,954,980]" italics="true" pageId="83" pageNumber="84">Edwards et al., 2017</emphasis>
|
||
</bibRefCitation>
|
||
). Therefore, should size dimorphism be found in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1065,1134,995,1020]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1065,1134,995,1020]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, it might also be present, ontogenetically, from start to finish. Therefore, the absence of a basal bifurcation in the ontogram shows that early size-independent dimorphism was absent from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1383,1452,1074,1099]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1383,1452,1074,1099]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,233,1419]" pageId="83" pageNumber="84">
|
||
Among crocodylians, size-independent sexual dimorphism is limited to the shape of the external naris in
|
||
<emphasis box="[699,920,1153,1179]" italics="true" pageId="83" pageNumber="84">Gavialis gangeticus</emphasis>
|
||
and
|
||
<taxonomicName baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[981,1192,1153,1180]" class="Reptilia" family="Tyrannosauridae" genus="Caiman" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="latirostris">
|
||
<emphasis box="[981,1192,1153,1180]" italics="true" pageId="83" pageNumber="84">Caiman latirostris</emphasis>
|
||
</taxonomicName>
|
||
and to the width of the interorbital end of the temporal roof in
|
||
<taxonomicName authorityName="Schneider" authorityYear="1801" box="[970,1192,1193,1219]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="porosus">
|
||
<emphasis box="[970,1192,1193,1219]" italics="true" pageId="83" pageNumber="84">Crocodylus porosus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[1211,1526,1193,1220]" journalOrPublisher="Acta Zoologica" pageId="83" pageNumber="84" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[1211,1526,1193,1220]" italics="true" pageId="83" pageNumber="84">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
). In contrast, variation in the shape of the bony naris and the interorbital region in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1449,1516,1234,1259]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1449,1516,1234,1259]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is ontogenetically controlled and dimorphism is not seen. In summary,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1317,1386,1274,1299]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1317,1386,1274,1299]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
</taxonomicName>
|
||
shows the pattern of variation (high ontogenetic variation, dimorphism absent, high individual variation) that is expected where only species recognition is at work (
|
||
<bibRefCitation author="Padian K & Horner JR" journalOrPublisher="Journal of Zoology" pageId="83" pageNumber="84" pagination="23 - 27" part="283" refId="ref55513" refString="Padian K, Horner JR. 2011. The definition of sexual selection and its implications for dinosaurian biology. Journal of Zoology 283 (1): 23 - 27 DOI 10.1111 / j. 1469 - 7998.2010.00761. x." title="The definition of sexual selection and its implications for dinosaurian biology" type="journal article" year="2011">
|
||
<emphasis italics="true" pageId="83" pageNumber="84">Padian & Horner, 2011</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,1458,1927]" box="[498,812,1458,1486]" pageId="83" pageNumber="84">
|
||
<heading bold="true" box="[498,812,1458,1486]" fontSize="11" level="3" pageId="83" pageNumber="84" reason="6">
|
||
<emphasis bold="true" box="[498,812,1458,1486]" italics="true" pageId="83" pageNumber="84">Size, maturity, and age</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,1458,1927]" pageId="83" pageNumber="84">
|
||
The suggestion has recently been made that size might correlate with maturity in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,982,1541,1568]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,1542,1567]" italics="true" pageId="83" pageNumber="84">T. rex</emphasis>
|
||
(
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[586,971,1541,1568]" journalOrPublisher="Anatomical Record" pageId="83" pageNumber="84" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[586,971,1541,1568]" italics="true" pageId="83" pageNumber="84">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, which was based on histological evidence. The specimen in question (
|
||
<materialsCitation box="[819,969,1581,1608]" collectionCode="RSM" pageId="83" pageNumber="84" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
), the youngest adult identified here, was hypothesized to exceed
|
||
<materialsCitation box="[772,954,1621,1647]" collectionCode="FMNH" pageId="83" pageNumber="84" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
in maturity based on the presence of a high degree of remodeling in the fibula that has produced a dense system of secondary osteons (
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" box="[508,894,1701,1727]" journalOrPublisher="Anatomical Record" pageId="83" pageNumber="84" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[508,894,1701,1727]" italics="true" pageId="83" pageNumber="84">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
). It is expected that if remodeling is congruent with maturity, then on the ontogram
|
||
<materialsCitation box="[875,1016,1740,1767]" collectionCode="RSM" pageId="83" pageNumber="84" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
would share a most recent growth stage with
|
||
<materialsCitation box="[498,683,1781,1807]" collectionCode="FMNH" pageId="83" pageNumber="84" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
to the exclusion of all other specimens.
|
||
</paragraph>
|
||
<paragraph blockId="83.[498,1542,1458,1927]" lastBlockId="84.[498,1542,234,1616]" lastPageId="84" lastPageNumber="85" pageId="83" pageNumber="84">
|
||
Among living archosaurs (e.g.,
|
||
<emphasis box="[894,1124,1820,1846]" italics="true" pageId="83" pageNumber="84">Melanosuchus niger</emphasis>
|
||
,
|
||
<emphasis box="[1137,1312,1821,1847]" italics="true" pageId="83" pageNumber="84">
|
||
Caiman
|
||
<taxonomicName authorityName="Daudin" authorityYear="1802" box="[1238,1312,1821,1846]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="83" pageNumber="84" phylum="Chordata" rank="species" species="yacare">yacare</taxonomicName>
|
||
</emphasis>
|
||
) variation in size is high, which is evidenced by high standard deviations (e.g.,
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[1190,1380,1860,1886]" journalOrPublisher="Salamandra" pageId="83" pageNumber="84" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[1190,1380,1860,1886]" italics="true" pageId="83" pageNumber="84">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Taylor P & Li F & Holland A & Martin M & Rosenblatt AE" journalOrPublisher="Amphibia-Reptilia" pageId="83" pageNumber="84" pagination="9 - 14" part="27" refId="ref56224" refString="Taylor P, Li F, Holland A, Martin M, Rosenblatt AE. 2016. Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana. Amphibia-Reptilia 27 (1): 9 - 14 DOI 10.1163 / 15685381 - 00003024." title="Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana" type="journal article" year="2016">
|
||
<emphasis italics="true" pageId="83" pageNumber="84">Taylor et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
), wide 95% confidence intervals (
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[953,1182,1900,1926]" journalOrPublisher="PLOS ONE" pageId="83" pageNumber="84" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[953,1182,1900,1926]" italics="true" pageId="83" pageNumber="84">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
), individual variation in growth rates (
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[663,891,234,261]" journalOrPublisher="PLOS ONE" pageId="84" pageNumber="85" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[663,891,234,261]" italics="true" pageId="84" pageNumber="85">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
), and by the relative inaccuracy of age estimation from size (
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[560,789,274,301]" journalOrPublisher="PLOS ONE" pageId="84" pageNumber="85" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[560,789,274,301]" italics="true" pageId="84" pageNumber="85">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
). Individual variation tends to be high among large, mature animals (e.g.,
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[660,768,314,340]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="acutus">
|
||
<emphasis box="[660,768,314,340]" italics="true" pageId="84" pageNumber="85">C. acutus</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[782,972,314,340]" journalOrPublisher="Salamandra" pageId="84" pageNumber="85" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[782,972,314,340]" italics="true" pageId="84" pageNumber="85">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
). Also, agreement between growth rate data and growth curve models are dependent upon sample size (
|
||
<bibRefCitation author="Taylor P & Li F & Holland A & Martin M & Rosenblatt AE" box="[1150,1362,354,380]" journalOrPublisher="Amphibia-Reptilia" pageId="84" pageNumber="85" pagination="9 - 14" part="27" refId="ref56224" refString="Taylor P, Li F, Holland A, Martin M, Rosenblatt AE. 2016. Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana. Amphibia-Reptilia 27 (1): 9 - 14 DOI 10.1163 / 15685381 - 00003024." title="Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana" type="journal article" year="2016">
|
||
<emphasis box="[1150,1362,354,380]" italics="true" pageId="84" pageNumber="85">Taylor et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
). This suggests that currently the sample size for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[898,966,394,419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[898,966,394,419]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(seven specimens in
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1219,1466,394,420]" journalOrPublisher="Nature" pageId="84" pageNumber="85" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1219,1466,394,420]" italics="true" pageId="84" pageNumber="85">Erickson et al. (2004)</emphasis>
|
||
</bibRefCitation>
|
||
) is insufficient for accurately capturing the congruence between mass and age.
|
||
</paragraph>
|
||
<paragraph blockId="84.[498,1542,234,1616]" pageId="84" pageNumber="85">
|
||
Also, size in crocodylians is affected by multiple extrinsic variables, including seasonality, seasonality of growth, incubation conditions, mean temperature (although this effect is not always found;
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[814,1043,553,579]" journalOrPublisher="PLOS ONE" pageId="84" pageNumber="85" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[814,1043,553,579]" italics="true" pageId="84" pageNumber="85">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
), rainfall, prey type, prey abundance, population density, social milieu, and disease (
|
||
<bibRefCitation author="Campos Z & Mourao G & Coutinho M & Magnusson WE & Rogers S." box="[1048,1277,593,619]" journalOrPublisher="PLOS ONE" pageId="84" pageNumber="85" pagination="e 89363" part="9" refId="ref52557" refString="Campos Z, Mourao G, Coutinho M, Magnusson WE, Rogers S. 2014. Growth of Caiman crocodilus yacare in the Brazilian Pantanal. PLOS ONE 9 (2): e 89363 DOI 10.1371 / journal. pone. 0089363." title="Growth of Caiman crocodilus yacare in the Brazilian Pantanal" type="journal article" year="2014">
|
||
<emphasis box="[1048,1277,593,619]" italics="true" pageId="84" pageNumber="85">Campos et al., 2014</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Taylor P & Li F & Holland A & Martin M & Rosenblatt AE" box="[1292,1504,593,619]" journalOrPublisher="Amphibia-Reptilia" pageId="84" pageNumber="85" pagination="9 - 14" part="27" refId="ref56224" refString="Taylor P, Li F, Holland A, Martin M, Rosenblatt AE. 2016. Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana. Amphibia-Reptilia 27 (1): 9 - 14 DOI 10.1163 / 15685381 - 00003024." title="Growth rates of black caiman (Melanosuchus niger) in the Rupununi region of Guyana" type="journal article" year="2016">
|
||
<emphasis box="[1292,1504,593,619]" italics="true" pageId="84" pageNumber="85">Taylor et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
). Although not all of these variables would have affected
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1134,1200,633,658]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1134,1200,633,658]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, an endothermic dinosaur, to the same extent that is seen in ectothermic crocodylians, it is not surprising that the sizes seen among the most mature specimens are not congruent with maturity. Alternatively, if extrinsic factors had a negligible effect upon body size, then sexual dimorphism might explain the noncongruence between maturity and size (but as shown, it does not). Among living crocodylians, linear measurements of adult animals are more variable than what is seen in juveniles (
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[965,1156,872,898]" journalOrPublisher="Salamandra" pageId="84" pageNumber="85" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[965,1156,872,898]" italics="true" pageId="84" pageNumber="85">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
), and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1233,1302,873,898]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1233,1302,873,898]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is consistent with that pattern.
|
||
</paragraph>
|
||
<paragraph blockId="84.[498,1542,234,1616]" pageId="84" pageNumber="85">
|
||
In
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[564,775,952,978]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[564,775,952,978]" italics="true" pageId="84" pageNumber="85">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
, growth rate effectively ceases at less than half the maximum age (
|
||
<bibRefCitation author="Wilkinson PM & Rainwater TR & Woodward AR & Leone EH & Carter C." box="[508,763,991,1017]" journalOrPublisher="Copeia" pageId="84" pageNumber="85" pagination="843 - 852" part="104" refId="ref56518" refString="Wilkinson PM, Rainwater TR, Woodward AR, Leone EH, Carter C. 2016. Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures. Copeia 104 (4): 843 - 852 DOI 10.1643 / CH- 16 - 430." title="Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures" type="journal article" year="2016">
|
||
<emphasis box="[508,763,991,1017]" italics="true" pageId="84" pageNumber="85">Wilkinson et al., 2016</emphasis>
|
||
</bibRefCitation>
|
||
); a similar pattern is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1110,1179,992,1017]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1110,1179,992,1017]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, where growth rate is greatly reduced at slightly greater than half the maximum age (18
|
||
<emphasis box="[1186,1201,1032,1057]" italics="true" pageId="84" pageNumber="85">–</emphasis>
|
||
28 years;
|
||
<bibRefCitation author="Horner JR & Padian K." journalOrPublisher="Proceedings of the Royal Society of London. Series B: Biological Sciences" pageId="84" pageNumber="85" pagination="1875 - 1880" part="271" refId="ref54442" refString="Horner JR, Padian K. 2004. Age and growth dynamics of Tyrannosaurus rex. Proceedings of the Royal Society of London. Series B: Biological Sciences 271 (1551): 1875 - 1880 DOI 10.1098 / rspb. 2004.2829." title="Age and growth dynamics of Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis italics="true" pageId="84" pageNumber="85">Horner & Padian, 2004</emphasis>
|
||
</bibRefCitation>
|
||
). The relatively wide range of size and mass for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1118,1185,1072,1097]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1118,1185,1072,1097]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA" box="[1203,1434,1071,1097]" journalOrPublisher="Nature" pageId="84" pageNumber="85" pagination="772 - 775" part="772 - 775" refId="ref53207" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 772 - 775 (7001): 772 - 775 DOI 10.1038 / nature 02699." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">
|
||
<emphasis box="[1203,1434,1071,1097]" italics="true" pageId="84" pageNumber="85">Erickson et al., 2004</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Persons WS & Currie PJ & Erickson GM" journalOrPublisher="Anatomical Record" pageId="84" pageNumber="85" pagination="656 - 672" part="303" refId="ref55578" refString="Persons WS, Currie PJ, Erickson GM. 2019. An older and exceptionally large adult specimen of Tyrannosaurus rex. Anatomical Record 303 (4): 656 - 672 DOI 10.1002 / ar. 24118." title="An older and exceptionally large adult specimen of Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis italics="true" pageId="84" pageNumber="85">Persons, Currie & Erickson, 2019</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Snively E & O ' Brien H & Henderson DM & Mallison H & Surring LA & Burns ME & Holtz TR Jr & Russell AP & Witmer LM & Currie PJ & Hartman SA & Cotton JR" box="[796,1013,1111,1137]" journalOrPublisher="PeerJ" pageId="84" pageNumber="85" pagination="e 6432" part="7" refId="ref56122" refString="Snively E, O ' Brien H, Henderson DM, Mallison H, Surring LA, Burns ME, Holtz TR Jr, Russell AP, Witmer LM, Currie PJ, Hartman SA, Cotton JR. 2019. Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods. PeerJ 7 (7447): e 6432 DOI 10.7717 / peerj. 6432." title="Lower rotational inertia and larger leg muscles indicate more rapid turns in tyrannosaurids than in other large theropods" type="journal article" year="2019">
|
||
<emphasis box="[796,1013,1111,1137]" italics="true" pageId="84" pageNumber="85">Snively et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
;) has two possible explanations: (1) it had determinate growth type II (sensu
|
||
<bibRefCitation author="Sebens KP" box="[908,1054,1151,1177]" journalOrPublisher="Annual Review of Ecology and Systematics" pageId="84" pageNumber="85" pagination="371 - 407" part="18" refId="ref56012" refString="Sebens KP. 1987. The ecology of indeterminate growth in animals. Annual Review of Ecology and Systematics 18 (1): 371 - 407 DOI 10.1146 / annurev. es. 18.110187.002103." title="The ecology of indeterminate growth in animals" type="journal article" year="1987">
|
||
<emphasis box="[908,1054,1151,1177]" italics="true" pageId="84" pageNumber="85">Sebens, 1987</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Wilkinson PM & Rainwater TR & Woodward AR & Leone EH & Carter C." box="[1095,1362,1151,1177]" journalOrPublisher="Copeia" pageId="84" pageNumber="85" pagination="843 - 852" part="104" refId="ref56518" refString="Wilkinson PM, Rainwater TR, Woodward AR, Leone EH, Carter C. 2016. Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures. Copeia 104 (4): 843 - 852 DOI 10.1643 / CH- 16 - 430." title="Determinate growth and reproductive lifespan in the American alligator (Alligator mississippiensis): evidence from long-term recaptures" type="journal article" year="2016">
|
||
<emphasis box="[1095,1362,1151,1177]" italics="true" pageId="84" pageNumber="85">Wilkinson et al. (2016)</emphasis>
|
||
</bibRefCitation>
|
||
) where although final size is genetically controlled it is affected by extrinsic factors, resulting in a wide range of adult sizes; alternatively (2) it had determinate growth type I where it has sexual size dimorphism, resulting in the scatter of adult sizes, but the environment has little effect upon asymptotic size. Therefore, the evidence shows that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1313,1381,1311,1336]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1313,1381,1311,1336]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
had determinate type II growth.
|
||
</paragraph>
|
||
<paragraph blockId="84.[498,1542,234,1616]" pageId="84" pageNumber="85">
|
||
The variation in size among the categories of adult (i.e., the noncongruence between size and maturity) is evidence of determinate growth, where the greatest amount of variation in size is seen that reflects the idiosyncratic differences in accumulated growth rate between individuals. In other words, the predictive value of size in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1333,1401,1510,1535]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1333,1401,1510,1535]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
ceases once the individual asymptote is reached (
|
||
<bibRefCitation author="Monteiro LR & Cavalcanti MJ & Sommer HJS III." box="[930,1378,1549,1576]" journalOrPublisher="Journal of Morphology" pageId="84" pageNumber="85" pagination="53 - 62" part="231" refId="ref55265" refString="Monteiro LR, Cavalcanti MJ, Sommer HJS III. 1997. Comparative ontogenetic shape changes in the skull of Caiman species (Crocodylia, Alligatoridae). Journal of Morphology 231: 53 - 62 DOI 10.1002 / (SICI) 1097 - 4687 (199701) 231: 1 <53 :: AID-JMOR 5> 3.0. CO; 2 - P." title="Comparative ontogenetic shape changes in the skull of Caiman species (Crocodylia, Alligatoridae)" type="journal article" year="1997">
|
||
<emphasis box="[930,1378,1549,1576]" italics="true" pageId="84" pageNumber="85">Monteiro, Cavalcanti & Sommer, 1997</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Tucker AD & Limpus CJ & McDonald KR & McCallum HI" journalOrPublisher="Queensland Australian Journal of Zoology" pageId="84" pageNumber="85" pagination="409 - 415" part="54" refId="ref56319" refString="Tucker AD, Limpus CJ, McDonald KR, McCallum HI. 2006. Growth dynamics of freshwater crocodiles (Crocodylus johnstoni) in the Lynd River. Queensland Australian Journal of Zoology 54 (6): 409 - 415 DOI 10.1071 / ZO 06099." title="Growth dynamics of freshwater crocodiles (Crocodylus johnstoni) in the Lynd River" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="84" pageNumber="85">Tucker et al., 2006</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="84.[498,1542,1660,1892]" box="[498,1205,1660,1690]" pageId="84" pageNumber="85">
|
||
<heading bold="true" box="[498,1205,1660,1690]" fontSize="12" level="2" pageId="84" pageNumber="85" reason="0">
|
||
<emphasis bold="true" box="[498,1205,1660,1690]" pageId="84" pageNumber="85">Plesiomorphic archosauriform growth patterns</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="84.[498,1542,1660,1892]" pageId="84" pageNumber="85">
|
||
A literature review of craniomandibular ontogeny in extinct (
|
||
<bibRefCitation author="Bhullar B-AS & Marugan-Lobon JM & Racimo F & Bever GS & Rowe TB & Norell MA & Abzhanov A." box="[1224,1446,1706,1732]" journalOrPublisher="Nature" pageId="84" pageNumber="85" pagination="223 - 226" part="487" refId="ref52184" refString="Bhullar B-AS, Marugan-Lobon JM, Racimo F, Bever GS, Rowe TB, Norell MA, Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. Nature 487 (7406): 223 - 226 DOI 10.1038 / nature 11146." title="Birds have paedomorphic dinosaur skulls" type="journal article" year="2012">
|
||
<emphasis box="[1224,1446,1706,1732]" italics="true" pageId="84" pageNumber="85">Bhullar et al., 2012</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[498,767,1746,1772]" journalOrPublisher="Journal of Anatomy" pageId="84" pageNumber="85" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[498,767,1746,1772]" italics="true" pageId="84" pageNumber="85">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[781,1153,1746,1772]" journalOrPublisher="PeerJ" pageId="84" pageNumber="85" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[781,1153,1746,1772]" italics="true" pageId="84" pageNumber="85">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[1167,1463,1746,1772]" journalOrPublisher="Proceedings of the Royal Society of London: Biology" pageId="84" pageNumber="85" pagination="2757 - 2761" part="273" refId="ref54486" refString="Horner JR, Goodwin MB. 2006. Major cranial changes during Triceratops ontogeny. Proceedings of the Royal Society of London: Biology 273 (1602): 2757 - 2761 DOI 10.1098 / rspb. 2006.3643." title="Major cranial changes during Triceratops ontogeny" type="journal article" year="2006">
|
||
<emphasis box="[1167,1463,1746,1772]" italics="true" pageId="84" pageNumber="85">Horner & Goodwin, 2006</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[1477,1532,1746,1772]" journalOrPublisher="PLOS ONE" pageId="84" pageNumber="85" pagination="e 7626" part="4" refId="ref54525" refString="Horner JR, Goodwin MB. 2009. Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLOS ONE 4 (10): e 7626 DOI 10.1371 / journal. pone. 0007626." title="Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus" type="journal article" year="2009">
|
||
<emphasis box="[1477,1532,1746,1772]" italics="true" pageId="84" pageNumber="85">2009</emphasis>
|
||
</bibRefCitation>
|
||
) and extant (
|
||
<bibRefCitation author="Dodson P." box="[636,791,1786,1812]" journalOrPublisher="Zoology" pageId="84" pageNumber="85" pagination="315 - 355" part="1975" refId="ref53056" refString="Dodson P. 1975. Functional and ecological significance of relative growth in Alligator. Zoology 1975: 315 - 355." title="Functional and ecological significance of relative growth in Alligator" type="journal article" year="1975">
|
||
<emphasis box="[636,791,1786,1812]" italics="true" pageId="84" pageNumber="85">Dodson, 1975</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Piras P & Colangelo P & Adams DC & Buscalioni AD & Cubo J & Kotsakis T & Meloro C & Raia P." box="[803,990,1786,1812]" journalOrPublisher="Evolution & Development" pageId="84" pageNumber="85" pagination="568 - 579" part="12" refId="ref55617" refString="Piras P, Colangelo P, Adams DC, Buscalioni AD, Cubo J, Kotsakis T, Meloro C, Raia P. 2010. The Gavialis - Tomistoma debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships. Evolution & Development 12 (6): 568 - 579 DOI 10.1111 / j. 1525 - 142 X. 2010.00442. x." title="The Gavialis - Tomistoma debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships" type="journal article" year="2010">
|
||
<emphasis box="[803,990,1786,1812]" italics="true" pageId="84" pageNumber="85">Piras et al., 2010</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[1025,1340,1786,1812]" journalOrPublisher="Acta Zoologica" pageId="84" pageNumber="85" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[1025,1340,1786,1812]" italics="true" pageId="84" pageNumber="85">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[1353,1536,1786,1812]" journalOrPublisher="Salamandra" pageId="84" pageNumber="85" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[1353,1536,1786,1812]" italics="true" pageId="84" pageNumber="85">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Wu XB & Xue H & Wu LS & Zhu JL & Wang RP" box="[498,678,1825,1851]" journalOrPublisher="Animal Biodiversity and Conservation" pageId="84" pageNumber="85" pagination="65 - 71" part="29" refId="ref56842" refString="Wu XB, Xue H, Wu LS, Zhu JL, Wang RP. 2006. Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population. Animal Biodiversity and Conservation 29: 65 - 71." title="Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population" type="journal article" year="2006">
|
||
<emphasis box="[498,678,1825,1851]" italics="true" pageId="84" pageNumber="85">Wu et al., 2006</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[719,1046,1825,1852]" journalOrPublisher="Acta Zoologica" pageId="84" pageNumber="85" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[719,1046,1825,1852]" italics="true" pageId="84" pageNumber="85">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
) archosaurs was done to identify the ancestral growth trends inherited by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[935,1003,1866,1891]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="84" pageNumber="85" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[935,1003,1866,1891]" italics="true" pageId="84" pageNumber="85">T. rex</emphasis>
|
||
</taxonomicName>
|
||
from its archosauriform ancestor.
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,234,858]" pageId="85" pageNumber="86">
|
||
Evidence for growth trends established in the common ancestors Archosauriformes (i.e., the common ancestor of
|
||
<taxonomicName authority="Archosauria" authorityName="Archosauria" baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[853,1014,274,300]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[853,1014,274,300]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
+ Archosauria, and all of its descendants, living and extinct), Archosauria (i.e., the common ancestor of
|
||
<taxonomicName box="[1235,1362,314,340]" class="Reptilia" kingdom="Animalia" order="Crocodylia" pageId="85" pageNumber="86" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
|
||
+
|
||
<taxonomicName authorityName="Owen" authorityYear="1842" box="[1396,1527,314,340]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
, and all of its descendants, living and extinct),
|
||
<taxonomicName authorityName="Owen" authorityYear="1842" box="[1042,1171,354,380]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
(i.e., the common ancestor of
|
||
<taxonomicName baseAuthorityName="Horner & Goodwin" baseAuthorityYear="2006" box="[530,660,394,419]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[530,660,394,419]" italics="true" pageId="85" pageNumber="86">Triceratops</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[721,793,394,419]" italics="true" pageId="85" pageNumber="86">Passer</emphasis>
|
||
, and all of its descendants, living and extinct),
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[1352,1471,394,420]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="85" pageNumber="86" phylum="Chordata" rank="order">Saurischia</taxonomicName>
|
||
(i.e., the common ancestor of
|
||
<taxonomicName authorityName="Owen" authorityYear="1854" box="[853,1038,433,459]" class="Reptilia" family="Massospondylidae" genus="Massospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[853,1038,433,459]" italics="true" pageId="85" pageNumber="86">Massospondylus</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1073,1141,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1073,1141,434,459]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and all of its descendants, living and extinct), and Neotheropoda (i.e., the common ancestor of
|
||
<taxonomicName box="[1302,1435,473,499]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1302,1435,473,499]" italics="true" pageId="85" pageNumber="86">Coelophysis</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1468,1535,474,499]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1468,1535,474,499]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and all of its descendants, living and extinct), were drawn from comparisons between
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,567,554,579]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,554,579]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, a sample of saurischian dinosaurs (
|
||
<bibRefCitation author="Bhullar B-AS & Marugan-Lobon JM & Racimo F & Bever GS & Rowe TB & Norell MA & Abzhanov A." box="[996,1217,553,579]" journalOrPublisher="Nature" pageId="85" pageNumber="86" pagination="223 - 226" part="487" refId="ref52184" refString="Bhullar B-AS, Marugan-Lobon JM, Racimo F, Bever GS, Rowe TB, Norell MA, Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. Nature 487 (7406): 223 - 226 DOI 10.1038 / nature 11146." title="Birds have paedomorphic dinosaur skulls" type="journal article" year="2012">
|
||
<emphasis box="[996,1217,553,579]" italics="true" pageId="85" pageNumber="86">Bhullar et al., 2012</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" journalOrPublisher="PeerJ" pageId="85" pageNumber="86" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis italics="true" pageId="85" pageNumber="86">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
), extant crocodylians, and the extinct
|
||
<taxonomicName authority="(Ezcurra & Butler, 2015)" baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1007,1468,593,619]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1007,1168,593,619]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
(
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1187,1458,593,619]" journalOrPublisher="Journal of Anatomy" pageId="85" pageNumber="86" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1187,1458,593,619]" italics="true" pageId="85" pageNumber="86">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
; the comparisons are summarized in
|
||
<tableCitation box="[885,986,633,659]" captionStart="Table 23" captionStartId="86.[213,268,434,456]" captionTargetBox="[203,1863,535,1801]" captionText="Table 23 Summary of the character evidence for shared ontogenetic trends throughout the ontogeny of Archosauriformes. Rows correspond to major growth trends identified in the left hand column and, in the following cells, the osteological correlates of the trend for each taxon; columns correspond to taxa. The taxonomic distribution of the reduction of cephalic ornamentation is given in the text. emf, external mandibular fenestra. See text for details and sources; see Fig. 33 for an illustrated summary." httpUri="http://table.plazi.org/id/DF5C84E2FF92FFFEFF33FE3A3007FDBF" pageId="85" pageNumber="86" tableUuid="DF5C84E2FF92FFFEFF33FE3A3007FDBF">Table 23</tableCitation>
|
||
. The character states were optimized onto a topology that was reconstructed in MacClade (
|
||
<bibRefCitation author="Maddison WP & Maddison DR" box="[1052,1394,673,699]" pageId="85" pageNumber="86" publicationUrl="http://macclade.org" refId="ref55064" refString="Maddison WP, Maddison DR. 2005. MacClade 4: analysis of phylogeny and character evolution. Version 4.08 a. Available at http: // macclade. org." title="MacClade 4: analysis of phylogeny and character evolution. Version 4.08 a" type="url" year="2005">
|
||
<emphasis box="[1052,1394,673,699]" italics="true" pageId="85" pageNumber="86">Maddison & Maddison, 2005</emphasis>
|
||
</bibRefCitation>
|
||
) to identify ancestral states in the phylogenetic hierarchy. The informal terms
|
||
<emphasis box="[1276,1288,712,738]" italics="true" pageId="85" pageNumber="86">“</emphasis>
|
||
archosauriforms
|
||
<emphasis box="[1478,1490,712,738]" italics="true" pageId="85" pageNumber="86">”</emphasis>
|
||
and
|
||
<emphasis box="[498,510,752,778]" italics="true" pageId="85" pageNumber="86">“</emphasis>
|
||
archosaurs
|
||
<emphasis box="[635,647,752,778]" italics="true" pageId="85" pageNumber="86">”</emphasis>
|
||
are used in place of their formal names in this discussion; the name
|
||
<taxonomicName box="[498,758,792,818]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[498,758,792,818]" italics="true" pageId="85" pageNumber="86">Tyrannosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
is used in place of
|
||
<emphasis box="[987,1143,792,818]" italics="true" pageId="85" pageNumber="86">“Tarbosaurus</emphasis>
|
||
.
|
||
<emphasis box="[1150,1162,792,818]" italics="true" pageId="85" pageNumber="86">”</emphasis>
|
||
A summary of the phylogenetic framework, the taxa included in this comparison, and the results are shown in
|
||
<figureCitation box="[1432,1511,832,858]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="85" pageNumber="86">Fig. 33</figureCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,897,1725]" box="[498,1079,897,925]" pageId="85" pageNumber="86">
|
||
<heading bold="true" box="[498,1079,897,925]" fontSize="11" level="3" pageId="85" pageNumber="86" reason="6">
|
||
<emphasis bold="true" box="[498,1079,897,925]" italics="true" pageId="85" pageNumber="86">Dorsoventral deepening of the skull frame</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,897,1725]" pageId="85" pageNumber="86">
|
||
Ontogenetic deepening of the skull frame comes from several lines of evidence, including the dorsoventral height of skull regions, specific bones, and fenestrae. In the archosauriforms
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[695,912,1021,1046]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[695,912,1021,1046]" italics="true" pageId="85" pageNumber="86">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
, crocodylians and
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1127,1288,1021,1047]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1127,1288,1021,1047]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
, the horizontal ramus of the maxilla increases in dorsoventral height (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1062,1333,1060,1087]" journalOrPublisher="Journal of Anatomy" pageId="85" pageNumber="86" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1062,1333,1060,1087]" italics="true" pageId="85" pageNumber="86">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
) (
|
||
<figureCitation box="[1361,1441,1060,1086]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="85" pageNumber="86">Fig. 33</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,897,1725]" pageId="85" pageNumber="86">
|
||
Another correlate of increased skull height in archosauriforms is the decreased height of the postorbital that is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[851,920,1141,1166]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[851,920,1141,1166]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, where the contribution of the bone to skull height is reduced by the dorsoventral expansion of the jugal, and
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1192,1353,1180,1206]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1192,1353,1180,1206]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
, where its height relative to total skull length decreases (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1033,1302,1220,1246]" journalOrPublisher="Journal of Anatomy" pageId="85" pageNumber="86" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1033,1302,1220,1246]" italics="true" pageId="85" pageNumber="86">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
). A reduction in the length of the orbital fenestra is another correlate of skull deepening, which is seen in
|
||
<taxonomicName authority="(Bhullar et al., 2012)" baseAuthorityName="Bhullar" baseAuthorityYear="2012" box="[498,882,1299,1326]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[498,631,1299,1325]" italics="true" pageId="85" pageNumber="86">Coelophysis</emphasis>
|
||
(
|
||
<bibRefCitation author="Bhullar B-AS & Marugan-Lobon JM & Racimo F & Bever GS & Rowe TB & Norell MA & Abzhanov A." box="[650,873,1299,1326]" journalOrPublisher="Nature" pageId="85" pageNumber="86" pagination="223 - 226" part="487" refId="ref52184" refString="Bhullar B-AS, Marugan-Lobon JM, Racimo F, Bever GS, Rowe TB, Norell MA, Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. Nature 487 (7406): 223 - 226 DOI 10.1038 / nature 11146." title="Birds have paedomorphic dinosaur skulls" type="journal article" year="2012">
|
||
<emphasis box="[650,873,1299,1326]" italics="true" pageId="85" pageNumber="86">Bhullar et al., 2012</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[944,1012,1300,1325]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[944,1012,1300,1325]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,897,1725]" pageId="85" pageNumber="86">
|
||
In archosaurs, the snout and jugal of crocodylians increase in depth, as also occurs in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[531,599,1380,1405]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[531,599,1380,1405]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, in
|
||
<emphasis box="[647,877,1379,1405]" italics="true" pageId="85" pageNumber="86">Melanosuchus niger</emphasis>
|
||
(
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[896,1211,1379,1406]" journalOrPublisher="Acta Zoologica" pageId="85" pageNumber="86" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[896,1211,1379,1406]" italics="true" pageId="85" pageNumber="86">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
),
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[1236,1344,1380,1406]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="acutus">
|
||
<emphasis box="[1236,1344,1380,1406]" italics="true" pageId="85" pageNumber="86">C. acutus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1825" class="Reptilia" family="Tyrannosauridae" genus="Mecistops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="cataphractus">
|
||
<emphasis italics="true" pageId="85" pageNumber="86">Mecistops cataphractus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName baseAuthorityName="Muller" baseAuthorityYear="1838" box="[713,954,1419,1445]" class="Reptilia" family="Crocodylidae" genus="Tomistoma" kingdom="Animalia" order="Crocodylia" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="schlegelii">
|
||
<emphasis box="[713,954,1419,1445]" italics="true" pageId="85" pageNumber="86">Tomistoma schlegelii</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[972,1287,1419,1446]" journalOrPublisher="Acta Zoologica" pageId="85" pageNumber="86" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[972,1287,1419,1446]" italics="true" pageId="85" pageNumber="86">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
); likewise, a shallow snout is a feature shared between juvenile caimans (
|
||
<bibRefCitation author="Foth C & Fernandez Blanco MV & Bona P & Scheyer TM" box="[1114,1303,1459,1485]" journalOrPublisher="Journal of Morphology" pageId="85" pageNumber="86" pagination="259 - 273" part="279" refId="ref53510" refString="Foth C, Fernandez Blanco MV, Bona P, Scheyer TM. 2018. Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri. Journal of Morphology 279 (2): 259 - 273 DOI 10.1002 / jmor. 20769." title="Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri" type="journal article" year="2018">
|
||
<emphasis box="[1114,1303,1459,1485]" italics="true" pageId="85" pageNumber="86">Foth et al., 2018</emphasis>
|
||
</bibRefCitation>
|
||
) and
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1374,1442,1460,1485]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1374,1442,1460,1485]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. This change is one of several correlates of the ontogenetic increase in height of the entire skull frame. Analysis of morphometric data quantifies the deepening of the skull, which is seen in
|
||
<taxonomicName authorityName="Owen" authorityYear="1854" box="[590,775,1578,1604]" class="Reptilia" family="Massospondylidae" genus="Massospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[590,775,1578,1604]" italics="true" pageId="85" pageNumber="86">Massospondylus</emphasis>
|
||
</taxonomicName>
|
||
, the megalosaurid
|
||
<taxonomicName authority=", Allosaurus" authorityName="Allosaurus" box="[1001,1336,1578,1605]" class="Reptilia" family="Megalosauridae" genus="Dubreuillosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1001,1200,1578,1604]" italics="true" pageId="85" pageNumber="86">Dubreuillosaurus</emphasis>
|
||
,
|
||
<emphasis box="[1214,1336,1578,1604]" italics="true" pageId="85" pageNumber="86">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Foth, Hendrick & Ezcurra, 2016)" baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1404,1513,1578,1604]" italics="true" pageId="85" pageNumber="86">T. bataar</emphasis>
|
||
(
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[508,882,1618,1645]" journalOrPublisher="PeerJ" pageId="85" pageNumber="86" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[508,882,1618,1645]" italics="true" pageId="85" pageNumber="86">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Optimization of the trends of increase in snout height and skull height onto the phylogenetic topology shows that both are ancestral for Archosauriformes, a trend that is not seen in
|
||
<taxonomicName box="[1038,1171,1698,1724]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1038,1171,1698,1724]" italics="true" pageId="85" pageNumber="86">Coelophysis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[1190,1269,1698,1724]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="85" pageNumber="86">Fig. 33</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,1763,1913]" box="[498,961,1763,1791]" pageId="85" pageNumber="86">
|
||
<heading bold="true" box="[498,961,1763,1791]" fontSize="11" level="3" pageId="85" pageNumber="86" reason="6">
|
||
<emphasis bold="true" box="[498,961,1763,1791]" italics="true" pageId="85" pageNumber="86">Expansion of the adductor region</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="85.[498,1542,1763,1913]" lastBlockId="88.[498,1525,234,301]" lastPageId="88" lastPageNumber="89" pageId="85" pageNumber="86">
|
||
In the archosauriforms
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[774,841,1808,1833]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[774,841,1808,1833]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[855,1016,1807,1833]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[855,1016,1807,1833]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
, and all crocodylian taxa, the orbital fenestra is ontogenetically reduced in size, a correlate of adductor expansion (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" journalOrPublisher="Journal of Anatomy" pageId="85" pageNumber="86" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis italics="true" pageId="85" pageNumber="86">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[567,875,1887,1913]" journalOrPublisher="Acta Zoologica" pageId="85" pageNumber="86" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[567,875,1887,1913]" italics="true" pageId="85" pageNumber="86">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Fernandez Blanco MV & Bona P & Scheyer TM" box="[888,1072,1887,1913]" journalOrPublisher="Journal of Morphology" pageId="85" pageNumber="86" pagination="259 - 273" part="279" refId="ref53510" refString="Foth C, Fernandez Blanco MV, Bona P, Scheyer TM. 2018. Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri. Journal of Morphology 279 (2): 259 - 273 DOI 10.1002 / jmor. 20769." title="Cranial shape variation in jacarean caimanines (Crocodylia, Alligatoroidea) and its implications in the taxonomic status of extinct species: the case of Melanosuchus fisheri" type="journal article" year="2018">
|
||
<emphasis box="[888,1072,1887,1913]" italics="true" pageId="85" pageNumber="86">Foth et al., 2018</emphasis>
|
||
</bibRefCitation>
|
||
). In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1128,1195,1888,1913]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="85" pageNumber="86" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1128,1195,1888,1913]" italics="true" pageId="85" pageNumber="86">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1253,1414,1887,1913]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="85" pageNumber="86" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1253,1414,1887,1913]" italics="true" pageId="85" pageNumber="86">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
the orbital fenestra decreases in length and increases in height (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1120,1390,1565,1592]" journalOrPublisher="Journal of Anatomy" pageId="87" pageNumber="88" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1120,1390,1565,1592]" italics="true" pageId="87" pageNumber="88">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
). Ergo, this reduction in size is almost certainly the result of two processes, namely the rostrocaudal expansion of the adductor chamber and dorsoventral expansion of the skull frame. The adductor region, including the laterotemporal fenestra, increases in height in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,566,1725,1750]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="87" pageNumber="88" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,566,1725,1750]" italics="true" pageId="87" pageNumber="88">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="(Ezcurra & Butler, 2015)" baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[625,1083,1725,1751]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="87" pageNumber="88" phylum="Chordata" rank="genus">
|
||
<emphasis box="[625,786,1725,1751]" italics="true" pageId="87" pageNumber="88">Proterosuchus</emphasis>
|
||
(
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[805,1073,1725,1751]" journalOrPublisher="Journal of Anatomy" pageId="87" pageNumber="88" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[805,1073,1725,1751]" italics="true" pageId="87" pageNumber="88">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Geometric morphometric comparison shows that the adductor region is not expanded in
|
||
<taxonomicName authorityName="Owen" authorityYear="1854" box="[1103,1289,1764,1790]" class="Reptilia" family="Massospondylidae" genus="Massospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="87" pageNumber="88" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1103,1289,1764,1790]" italics="true" pageId="87" pageNumber="88">Massospondylus</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName box="[1331,1464,1764,1790]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="87" pageNumber="88" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1331,1464,1764,1790]" italics="true" pageId="87" pageNumber="88">Coelophysis</emphasis>
|
||
</taxonomicName>
|
||
, whereas it is expanded in
|
||
<taxonomicName authority=", Allosaurus" authorityName="Allosaurus" box="[806,1142,1804,1831]" class="Reptilia" family="Megalosauridae" genus="Dubreuillosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="87" pageNumber="88" phylum="Chordata" rank="genus">
|
||
<emphasis box="[806,1005,1804,1830]" italics="true" pageId="87" pageNumber="88">Dubreuillosaurus</emphasis>
|
||
,
|
||
<emphasis box="[1020,1142,1804,1830]" italics="true" pageId="87" pageNumber="88">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName authority="(Foth, Hendrick & Ezcurra, 2016)" baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="87" pageNumber="88" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1210,1318,1804,1830]" italics="true" pageId="87" pageNumber="88">T. bataar</emphasis>
|
||
(
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[508,880,1844,1871]" journalOrPublisher="PeerJ" pageId="87" pageNumber="88" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[508,880,1844,1871]" italics="true" pageId="87" pageNumber="88">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Optimization of character states onto the phylogenetic topology recovers adductor chamber expansion as plesiomorphic for Archosauriformes (
|
||
<figureCitation box="[508,588,274,300]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="88" pageNumber="89">Fig. 33</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FF92FFFEFF33FE3A3007FDBF" httpUri="http://table.plazi.org/id/DF5C84E2FF92FFFEFF33FE3A3007FDBF" pageId="86" pageNumber="87" startId="86.[213,268,434,456]" targetBox="[203,1863,535,1801]" targetIsTable="true" targetPageId="86">
|
||
<paragraph blockId="86.[213,1854,434,515]" pageId="86" pageNumber="87">
|
||
<emphasis bold="true" box="[213,1448,434,456]" pageId="86" pageNumber="87">Table 23 Summary of the character evidence for shared ontogenetic trends throughout the ontogeny of Archosauriformes.</emphasis>
|
||
Rows correspond to major growth trends identified in the left hand column and, in the following cells, the osteological correlates of the trend for each taxon; columns correspond to taxa. The taxonomic distribution of the reduction of cephalic ornamentation is given in the text. emf, external mandibular fenestra. See text for details and sources; see Fig. 33 for an illustrated summary.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="86" pageNumber="87">
|
||
<table box="[203,1863,535,1801]" gridcols="9" gridrows="13" pageId="86" pageNumber="87">
|
||
<tr box="[203,1863,535,587]" gridrow="0" pageId="86" pageNumber="87">
|
||
<th box="[203,340,535,587]" gridcol="0" gridrow="0" pageId="86" pageNumber="87">Character/ taxon</th>
|
||
<th box="[356,604,535,587]" gridcol="1" gridrow="0" pageId="86" pageNumber="87">
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[356,494,535,557]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="86" pageNumber="87" phylum="Chordata" rank="genus">
|
||
<emphasis bold="true" box="[356,494,535,557]" italics="true" pageId="86" pageNumber="87">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<taxonomicName authority="Massospondylus Coelophysis Dubreuillosaurus Allosaurus" authorityName="Massospondylus Coelophysis Dubreuillosaurus Allosaurus" box="[627,1460,535,557]" class="Reptilia" kingdom="Animalia" order="Crocodylia" pageId="86" pageNumber="87" phylum="Chordata" rank="order">
|
||
<th box="[627,850,535,587]" gridcol="2" gridrow="0" pageId="86" pageNumber="87">Crocodylia</th>
|
||
<th box="[866,1024,535,587]" gridcol="3" gridrow="0" pageId="86" pageNumber="87">Massospondylus</th>
|
||
<th box="[1040,1155,535,587]" gridcol="4" gridrow="0" pageId="86" pageNumber="87">Coelophysis</th>
|
||
<th box="[1170,1340,535,587]" gridcol="5" gridrow="0" pageId="86" pageNumber="87">Dubreuillosaurus</th>
|
||
<th box="[1356,1476,535,587]" gridcol="6" gridrow="0" pageId="86" pageNumber="87">Allosaurus</th>
|
||
</taxonomicName>
|
||
<th box="[1491,1642,535,587]" gridcol="7" gridrow="0" pageId="86" pageNumber="87">
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[1491,1640,535,587]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="86" pageNumber="87" phylum="Chordata" rank="species" species="bataar">
|
||
Tyrannosaurus
|
||
<emphasis bold="true" box="[1491,1556,565,587]" italics="true" pageId="86" pageNumber="87">bataar</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<th box="[1663,1863,535,587]" gridcol="8" gridrow="0" pageId="86" pageNumber="87">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1663,1722,535,557]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="86" pageNumber="87" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[1663,1722,535,557]" italics="true" pageId="86" pageNumber="87">T. rex</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
</tr>
|
||
<tr box="[203,1863,606,686]" gridrow="1" pageId="86" pageNumber="87">
|
||
<th box="[203,340,606,686]" gridcol="0" gridrow="1" pageId="86" pageNumber="87">Increase in snout height</th>
|
||
<td box="[356,604,606,686]" gridcol="1" gridrow="1" pageId="86" pageNumber="87">height maxillary horizontal ramus</td>
|
||
<td box="[627,850,606,686]" gridcol="2" gridrow="1" pageId="86" pageNumber="87">snout height</td>
|
||
<td box="[866,1024,606,686]" gridcol="3" gridrow="1" pageId="86" pageNumber="87">snout height</td>
|
||
<td box="[1040,1155,606,686]" gridcol="4" gridrow="1" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1170,1340,606,686]" gridcol="5" gridrow="1" pageId="86" pageNumber="87">snout height</td>
|
||
<td box="[1356,1476,606,686]" gridcol="6" gridrow="1" pageId="86" pageNumber="87">snout height</td>
|
||
<td box="[1491,1642,606,686]" gridcol="7" gridrow="1" pageId="86" pageNumber="87">snout height</td>
|
||
<td box="[1663,1863,606,686]" gridcol="8" gridrow="1" pageId="86" pageNumber="87">height maxillary horizontal ramus, snout height</td>
|
||
</tr>
|
||
<tr box="[203,1863,702,1016]" gridrow="2" pageId="86" pageNumber="87">
|
||
<th box="[203,340,702,1016]" gridcol="0" gridrow="2" pageId="86" pageNumber="87">Increase skull height</th>
|
||
<td box="[356,604,702,1016]" gridcol="1" gridrow="2" pageId="86" pageNumber="87">height orbital & laterotemporal fenestrae</td>
|
||
<td box="[627,850,702,1016]" gridcol="2" gridrow="2" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,702,1016]" gridcol="3" gridrow="2" pageId="86" pageNumber="87">enlarged postorbital, dorsal shift bony naris</td>
|
||
<td box="[1040,1155,702,1016]" gridcol="4" gridrow="2" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1170,1340,702,1016]" gridcol="5" gridrow="2" pageId="86" pageNumber="87">caudally inclined lacrimal, postorbital enlarged</td>
|
||
<td box="[1356,1476,702,1016]" gridcol="6" gridrow="2" pageId="86" pageNumber="87">lacrimal ventral ramus size, postorbital region height, angulation between maxilla and jugal</td>
|
||
<td box="[1491,1642,702,1016]" gridcol="7" gridrow="2" pageId="86" pageNumber="87">lacrimal ventral ramus size, postorbital region height, dorsal shift</td>
|
||
<td box="[1663,1863,702,1016]" gridcol="8" gridrow="2" pageId="86" pageNumber="87">height orbital & laterotemporal fenestrae, caudally inclined lacrimal, postorbital enlarged, postorbital region height, angulation between maxilla and jugal</td>
|
||
</tr>
|
||
<tr box="[203,1863,1032,1141]" gridrow="3" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1032,1141]" gridcol="0" gridrow="3" pageId="86" pageNumber="87">Increase in adductor chamber size</th>
|
||
<td box="[356,604,1032,1141]" gridcol="1" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[627,850,1032,1141]" gridcol="2" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[866,1024,1032,1141]" gridcol="3" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[1040,1155,1032,1141]" gridcol="4" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[1170,1340,1032,1141]" gridcol="5" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[1356,1476,1032,1141]" gridcol="6" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[1491,1642,1032,1141]" gridcol="7" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
<td box="[1663,1863,1032,1141]" gridcol="8" gridrow="3" pageId="86" pageNumber="87">length orbital fenestra</td>
|
||
</tr>
|
||
<tr box="[203,1863,1157,1208]" gridrow="4" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1157,1208]" gridcol="0" gridrow="4" pageId="86" pageNumber="87">Increase in skull width</th>
|
||
<td box="[356,604,1157,1208]" gridcol="1" gridrow="4" pageId="86" pageNumber="87">width frontoparietal region</td>
|
||
<td box="[627,850,1157,1208]" gridcol="2" gridrow="4" pageId="86" pageNumber="87">width frontoparietal region, width snout</td>
|
||
<td box="[866,1024,1157,1208]" gridcol="3" gridrow="4" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1157,1208]" gridcol="4" gridrow="4" pageId="86" pageNumber="87">width snout</td>
|
||
<td box="[1170,1340,1157,1208]" gridcol="5" gridrow="4" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1157,1208]" gridcol="6" gridrow="4" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1157,1208]" gridcol="7" gridrow="4" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1157,1208]" gridcol="8" gridrow="4" pageId="86" pageNumber="87">width frontoparietal region, width snout</td>
|
||
</tr>
|
||
<tr box="[203,1863,1224,1275]" gridrow="5" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1224,1275]" gridcol="0" gridrow="5" pageId="86" pageNumber="87">Antorbital fossa length</th>
|
||
<td box="[356,604,1224,1275]" gridcol="1" gridrow="5" pageId="86" pageNumber="87">?</td>
|
||
<td box="[627,850,1224,1275]" gridcol="2" gridrow="5" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1224,1275]" gridcol="3" gridrow="5" pageId="86" pageNumber="87">decrease</td>
|
||
<td box="[1040,1155,1224,1275]" gridcol="4" gridrow="5" pageId="86" pageNumber="87">increase</td>
|
||
<td box="[1170,1340,1224,1275]" gridcol="5" gridrow="5" pageId="86" pageNumber="87">decrease</td>
|
||
<td box="[1356,1476,1224,1275]" gridcol="6" gridrow="5" pageId="86" pageNumber="87">decrease</td>
|
||
<td box="[1491,1642,1224,1275]" gridcol="7" gridrow="5" pageId="86" pageNumber="87">decrease</td>
|
||
<td box="[1663,1863,1224,1275]" gridcol="8" gridrow="5" pageId="86" pageNumber="87">?</td>
|
||
</tr>
|
||
<tr box="[203,1863,1291,1371]" gridrow="6" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1291,1371]" gridcol="0" gridrow="6" pageId="86" pageNumber="87">Rostrocaudal jaw joint position</th>
|
||
<td box="[356,604,1291,1371]" gridcol="1" gridrow="6" pageId="86" pageNumber="87">caudal</td>
|
||
<td box="[627,850,1291,1371]" gridcol="2" gridrow="6" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1291,1371]" gridcol="3" gridrow="6" pageId="86" pageNumber="87">rostral</td>
|
||
<td box="[1040,1155,1291,1371]" gridcol="4" gridrow="6" pageId="86" pageNumber="87">caudal</td>
|
||
<td box="[1170,1340,1291,1371]" gridcol="5" gridrow="6" pageId="86" pageNumber="87">caudal</td>
|
||
<td box="[1356,1476,1291,1371]" gridcol="6" gridrow="6" pageId="86" pageNumber="87">rostral</td>
|
||
<td box="[1491,1642,1291,1371]" gridcol="7" gridrow="6" pageId="86" pageNumber="87">caudal</td>
|
||
<td box="[1663,1863,1291,1371]" gridcol="8" gridrow="6" pageId="86" pageNumber="87">?</td>
|
||
</tr>
|
||
<tr box="[203,1863,1387,1467]" gridrow="7" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1387,1467]" gridcol="0" gridrow="7" pageId="86" pageNumber="87">Dorsoventral jaw joint position</th>
|
||
<td box="[356,604,1387,1467]" gridcol="1" gridrow="7" pageId="86" pageNumber="87">ventral</td>
|
||
<td box="[627,850,1387,1467]" gridcol="2" gridrow="7" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1387,1467]" gridcol="3" gridrow="7" pageId="86" pageNumber="87">ventral</td>
|
||
<td box="[1040,1155,1387,1467]" gridcol="4" gridrow="7" pageId="86" pageNumber="87">dorsal</td>
|
||
<td box="[1170,1340,1387,1467]" gridcol="5" gridrow="7" pageId="86" pageNumber="87">no change</td>
|
||
<td box="[1356,1476,1387,1467]" gridcol="6" gridrow="7" pageId="86" pageNumber="87">ventral</td>
|
||
<td box="[1491,1642,1387,1467]" gridcol="7" gridrow="7" pageId="86" pageNumber="87">ventral</td>
|
||
<td box="[1663,1863,1387,1467]" gridcol="8" gridrow="7" pageId="86" pageNumber="87">?</td>
|
||
</tr>
|
||
<tr box="[203,1863,1483,1563]" gridrow="8" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1483,1563]" gridcol="0" gridrow="8" pageId="86" pageNumber="87">Increase of mandible height</th>
|
||
<td box="[356,604,1483,1563]" gridcol="1" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[627,850,1483,1563]" gridcol="2" gridrow="8" pageId="86" pageNumber="87">height mandible & emf</td>
|
||
<td box="[866,1024,1483,1563]" gridcol="3" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1483,1563]" gridcol="4" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1170,1340,1483,1563]" gridcol="5" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1483,1563]" gridcol="6" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1483,1563]" gridcol="7" gridrow="8" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1483,1563]" gridcol="8" gridrow="8" pageId="86" pageNumber="87">height mandible & emf</td>
|
||
</tr>
|
||
<tr box="[203,1863,1579,1630]" gridrow="9" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1579,1630]" gridcol="0" gridrow="9" pageId="86" pageNumber="87">Tooth number</th>
|
||
<td box="[356,604,1579,1630]" gridcol="1" gridrow="9" pageId="86" pageNumber="87">increase maxillary teeth</td>
|
||
<td box="[627,850,1579,1630]" gridcol="2" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1579,1630]" gridcol="3" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1579,1630]" gridcol="4" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1170,1340,1579,1630]" gridcol="5" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1579,1630]" gridcol="6" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1579,1630]" gridcol="7" gridrow="9" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1579,1630]" gridcol="8" gridrow="9" pageId="86" pageNumber="87">increase maxillary teeth</td>
|
||
</tr>
|
||
<tr box="[203,1863,1645,1667]" gridrow="10" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1645,1667]" gridcol="0" gridrow="10" pageId="86" pageNumber="87">Bite force</th>
|
||
<td box="[356,604,1645,1667]" gridcol="1" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[627,850,1645,1667]" gridcol="2" gridrow="10" pageId="86" pageNumber="87">increase</td>
|
||
<td box="[866,1024,1645,1667]" gridcol="3" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1645,1667]" gridcol="4" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1170,1340,1645,1667]" gridcol="5" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1645,1667]" gridcol="6" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1645,1667]" gridcol="7" gridrow="10" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1645,1667]" gridcol="8" gridrow="10" pageId="86" pageNumber="87">increase</td>
|
||
</tr>
|
||
<tr box="[203,1863,1683,1734]" gridrow="11" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1683,1734]" gridcol="0" gridrow="11" pageId="86" pageNumber="87">Deltopectoral crest</th>
|
||
<td box="[356,604,1683,1734]" gridcol="1" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[627,850,1683,1734]" gridcol="2" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1683,1734]" gridcol="3" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1683,1734]" gridcol="4" gridrow="11" pageId="86" pageNumber="87">enlarged crest</td>
|
||
<td box="[1170,1340,1683,1734]" gridcol="5" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1683,1734]" gridcol="6" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1683,1734]" gridcol="7" gridrow="11" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1683,1734]" gridcol="8" gridrow="11" pageId="86" pageNumber="87">enlarged crest</td>
|
||
</tr>
|
||
<tr box="[203,1863,1750,1801]" gridrow="12" pageId="86" pageNumber="87">
|
||
<th box="[203,340,1750,1801]" gridcol="0" gridrow="12" pageId="86" pageNumber="87">Trochanteric crest</th>
|
||
<td box="[356,604,1750,1801]" gridcol="1" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[627,850,1750,1801]" gridcol="2" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[866,1024,1750,1801]" gridcol="3" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1040,1155,1750,1801]" gridcol="4" gridrow="12" pageId="86" pageNumber="87">enhanced crest</td>
|
||
<td box="[1170,1340,1750,1801]" gridcol="5" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1356,1476,1750,1801]" gridcol="6" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1491,1642,1750,1801]" gridcol="7" gridrow="12" pageId="86" pageNumber="87">?</td>
|
||
<td box="[1663,1863,1750,1801]" gridcol="8" gridrow="12" pageId="86" pageNumber="87">enhanced crest</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.5825040" ID-Zenodo-Dep="5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="87" pageNumber="88" startId="87.[524,589,1254,1276]" targetBox="[498,1542,234,1242]" targetPageId="87">
|
||
<paragraph blockId="87.[524,1515,1254,1510]" pageId="87" pageNumber="88">
|
||
<emphasis bold="true" pageId="87" pageNumber="88">Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies.</emphasis>
|
||
Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1335,1515,1401,1422]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="87" pageNumber="88" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1335,1515,1401,1422]" italics="true" pageId="87" pageNumber="88">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa.
|
||
</paragraph>
|
||
<paragraph blockId="87.[524,1515,1254,1510]" box="[1099,1515,1488,1510]" pageId="87" pageNumber="88">Full-size DOI: 10.7717/peerj.9192/fig-33</paragraph>
|
||
</caption>
|
||
<paragraph blockId="88.[498,1541,337,1124]" box="[498,1050,337,365]" pageId="88" pageNumber="89">
|
||
<heading bold="true" box="[498,1050,337,365]" fontSize="11" level="3" pageId="88" pageNumber="89" reason="6">
|
||
<emphasis bold="true" box="[498,1050,337,365]" italics="true" pageId="88" pageNumber="89">Mediolateral widening of the skull frame</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1541,337,1124]" pageId="88" pageNumber="89">
|
||
In archosauriforms, an overall increase in snout width is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1266,1334,381,406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1266,1334,381,406]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and in the crocodylians
|
||
<taxonomicName authorityName="Fauvel" authorityYear="1879" box="[654,851,420,446]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="sinensis">
|
||
<emphasis box="[654,851,420,446]" italics="true" pageId="88" pageNumber="89">Alligator sinensis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Wu XB & Xue H & Wu LS & Zhu JL & Wang RP" box="[870,1049,420,446]" journalOrPublisher="Animal Biodiversity and Conservation" pageId="88" pageNumber="89" pagination="65 - 71" part="29" refId="ref56842" refString="Wu XB, Xue H, Wu LS, Zhu JL, Wang RP. 2006. Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population. Animal Biodiversity and Conservation 29: 65 - 71." title="Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population" type="journal article" year="2006">
|
||
<emphasis box="[870,1049,420,446]" italics="true" pageId="88" pageNumber="89">Wu et al., 2006</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[1090,1417,420,447]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[1090,1417,420,447]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
),
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[498,709,460,486]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="acutus">
|
||
<emphasis box="[498,709,460,486]" italics="true" pageId="88" pageNumber="89">Crocodylus acutus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1825" box="[724,992,460,486]" class="Reptilia" family="Tyrannosauridae" genus="Mecistops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="cataphractus">
|
||
<emphasis box="[724,992,460,486]" italics="true" pageId="88" pageNumber="89">Mecistops cataphractus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName baseAuthorityName="Muller" baseAuthorityYear="1838" box="[1059,1300,460,486]" class="Reptilia" family="Crocodylidae" genus="Tomistoma" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="schlegelii">
|
||
<emphasis box="[1059,1300,460,486]" italics="true" pageId="88" pageNumber="89">Tomistoma schlegelii</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Piras P & Colangelo P & Adams DC & Buscalioni AD & Cubo J & Kotsakis T & Meloro C & Raia P." box="[1319,1514,460,486]" journalOrPublisher="Evolution & Development" pageId="88" pageNumber="89" pagination="568 - 579" part="12" refId="ref55617" refString="Piras P, Colangelo P, Adams DC, Buscalioni AD, Cubo J, Kotsakis T, Meloro C, Raia P. 2010. The Gavialis - Tomistoma debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships. Evolution & Development 12 (6): 568 - 579 DOI 10.1111 / j. 1525 - 142 X. 2010.00442. x." title="The Gavialis - Tomistoma debate: the contribution of skull ontogenetic allometry and growth trajectories to the study of crocodylian relationships" type="journal article" year="2010">
|
||
<emphasis box="[1319,1514,460,486]" italics="true" pageId="88" pageNumber="89">Piras et al., 2010</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[530,854,500,526]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[530,854,500,526]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[869,1059,500,526]" journalOrPublisher="Salamandra" pageId="88" pageNumber="89" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[869,1059,500,526]" italics="true" pageId="88" pageNumber="89">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
), and also in
|
||
<taxonomicName baseAuthorityName="Hurlburt, Ridgley & Witmer" baseAuthorityYear="2013" box="[1220,1430,500,526]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis box="[1220,1430,500,526]" italics="true" pageId="88" pageNumber="89">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Dodson P." journalOrPublisher="Zoology" pageId="88" pageNumber="89" pagination="315 - 355" part="1975" refId="ref53056" refString="Dodson P. 1975. Functional and ecological significance of relative growth in Alligator. Zoology 1975: 315 - 355." title="Functional and ecological significance of relative growth in Alligator" type="journal article" year="1975">
|
||
<emphasis italics="true" pageId="88" pageNumber="89">Dodson, 1975</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[594,921,540,566]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[594,921,540,566]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
) and
|
||
<taxonomicName authorityName="Schmidt" authorityYear="1928" box="[993,1182,540,566]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="novaeguineae">
|
||
<emphasis box="[993,1182,540,566]" italics="true" pageId="88" pageNumber="89">C. novaeguineae</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Hall PM & Portier KM" box="[1201,1441,540,566]" journalOrPublisher="Herpetological Monographs" pageId="88" pageNumber="89" pagination="203 - 225" part="8" refId="ref54030" refString="Hall PM, Portier KM. 1994. Cranial morphometry of New Guinea crocodiles (Crocodylus novaeguineae): ontogenetic variation in relative growth of the skull and an assessment of its utility as a predictor of sex and size of individuals. Herpetological Monographs 8: 203 - 225 DOI 10.2307 / 1467081." title="Cranial morphometry of New Guinea crocodiles (Crocodylus novaeguineae): ontogenetic variation in relative growth of the skull and an assessment of its utility as a predictor of sex and size of individuals" type="journal article" year="1994">
|
||
<emphasis box="[1201,1441,540,566]" italics="true" pageId="88" pageNumber="89">Hall & Portier, 1994</emphasis>
|
||
</bibRefCitation>
|
||
in
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[498,817,580,606]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[498,817,580,606]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo (2013)</emphasis>
|
||
</bibRefCitation>
|
||
). Widening of the snout in crocodylians is thought to coincide with changes in diet, for example from invertebrates to vertebrates (
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[1302,1494,619,646]" journalOrPublisher="Salamandra" pageId="88" pageNumber="89" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[1302,1494,619,646]" italics="true" pageId="88" pageNumber="89">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1541,337,1124]" pageId="88" pageNumber="89">
|
||
An ontogenetic increase in frontal (=interorbital) width is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1313,1533,660,685]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1313,1533,660,685]" italics="true" pageId="88" pageNumber="89">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
and all crocodylians except
|
||
<taxonomicName authorityName="Fauvel" authorityYear="1879" box="[826,948,700,726]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="sinensis">
|
||
<emphasis box="[826,948,700,726]" italics="true" pageId="88" pageNumber="89">A. sinensis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Wu XB & Xue H & Wu LS & Zhu JL & Wang RP" box="[966,1146,699,725]" journalOrPublisher="Animal Biodiversity and Conservation" pageId="88" pageNumber="89" pagination="65 - 71" part="29" refId="ref56842" refString="Wu XB, Xue H, Wu LS, Zhu JL, Wang RP. 2006. Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population. Animal Biodiversity and Conservation 29: 65 - 71." title="Regression analysis between body and head measurements of Chinese alligators (Alligator sinensis) in the captive population" type="journal article" year="2006">
|
||
<emphasis box="[966,1146,699,725]" italics="true" pageId="88" pageNumber="89">Wu et al., 2006</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[1161,1476,699,726]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[1161,1476,699,726]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Edwards GP & Webb GJ & Manolis SC & Mazanov A." box="[498,734,739,765]" journalOrPublisher="Australian Journal of Zoology" pageId="88" pageNumber="89" pagination="97" part="65" refId="ref53078" refString="Edwards GP, Webb GJ, Manolis SC, Mazanov A. 2017. Morphometric analysis of the Australian freshwater crocodile (Crocodylus johnstoni). Australian Journal of Zoology 65 (2): 97 DOI 10.1071 / ZO 16079." title="Morphometric analysis of the Australian freshwater crocodile (Crocodylus johnstoni)" type="journal article" year="2017">
|
||
<emphasis box="[498,734,739,765]" italics="true" pageId="88" pageNumber="89">Edwards et al., 2017</emphasis>
|
||
</bibRefCitation>
|
||
). The ontogenetic widening of the parietals that is seen in
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[498,659,779,805]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[498,659,779,805]" italics="true" pageId="88" pageNumber="89">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
might be homologous to the increase in width seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1314,1383,780,805]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1314,1383,780,805]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and crocodylians (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[664,935,819,845]" journalOrPublisher="Journal of Anatomy" pageId="88" pageNumber="89" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[664,935,819,845]" italics="true" pageId="88" pageNumber="89">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1541,337,1124]" pageId="88" pageNumber="89">
|
||
Ontogenetic widening of the skull in archosaurs is indicated by correlates seen in the snout and dorsal skull roof. In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[866,935,899,924]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[866,935,899,924]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, caimans (e.g.,
|
||
<taxonomicName baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[1118,1260,898,925]" class="Reptilia" family="Tyrannosauridae" genus="Caiman" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="latirostris">
|
||
<emphasis box="[1118,1260,898,925]" italics="true" pageId="88" pageNumber="89">C. latirostris</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName box="[1276,1396,898,925]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="sclerops">
|
||
<emphasis box="[1276,1396,898,925]" italics="true" pageId="88" pageNumber="89">C. sclerops</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Daudin" authorityYear="1802" box="[1412,1519,899,925]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="yacare">
|
||
<emphasis box="[1412,1519,899,925]" italics="true" pageId="88" pageNumber="89">C. yacare</emphasis>
|
||
</taxonomicName>
|
||
), and crocodiles (e.g.,
|
||
<emphasis box="[740,960,938,965]" italics="true" pageId="88" pageNumber="89">
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[740,952,938,965]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="acutus">Crocodylus acutus</taxonomicName>
|
||
)
|
||
</emphasis>
|
||
the rostral end of the snout widens (Platt at al., 2011). The premaxilla shortens in
|
||
<emphasis box="[906,1136,978,1004]" italics="true" pageId="88" pageNumber="89">Melanosuchus niger</emphasis>
|
||
(
|
||
<bibRefCitation author="Foth C & Bona P & Desojo JB" box="[1155,1470,978,1005]" journalOrPublisher="Acta Zoologica" pageId="88" pageNumber="89" pagination="1 - 13" part="96" refId="ref53423" refString="Foth C, Bona P, Desojo JB. 2013. Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae). Acta Zoologica 96 (1): 1 - 13 DOI 10.1111 / azo. 12045." title="Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae, Caimanainae)" type="journal article" year="2013">
|
||
<emphasis box="[1155,1470,978,1005]" italics="true" pageId="88" pageNumber="89">Foth, Bona & Desojo, 2013</emphasis>
|
||
</bibRefCitation>
|
||
) and
|
||
<taxonomicName authority="(Bhullar et al., 2012)" baseAuthorityName="Bhullar" baseAuthorityYear="2012" box="[498,883,1018,1044]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[498,631,1018,1044]" italics="true" pageId="88" pageNumber="89">Coelophysis</emphasis>
|
||
(
|
||
<bibRefCitation author="Bhullar B-AS & Marugan-Lobon JM & Racimo F & Bever GS & Rowe TB & Norell MA & Abzhanov A." box="[650,873,1018,1044]" journalOrPublisher="Nature" pageId="88" pageNumber="89" pagination="223 - 226" part="487" refId="ref52184" refString="Bhullar B-AS, Marugan-Lobon JM, Racimo F, Bever GS, Rowe TB, Norell MA, Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. Nature 487 (7406): 223 - 226 DOI 10.1038 / nature 11146." title="Birds have paedomorphic dinosaur skulls" type="journal article" year="2012">
|
||
<emphasis box="[650,873,1018,1044]" italics="true" pageId="88" pageNumber="89">Bhullar et al., 2012</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, which corresponds to the widening and medialward reorientation of the premaxilla (i.e., increase in snout width) in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1252,1321,1059,1084]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1252,1321,1059,1084]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Optimization of skull widening on the topology recovers it as ancestral for Archosauriformes (
|
||
<figureCitation box="[1419,1498,1098,1124]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="88" pageNumber="89">Fig. 33</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1542,1160,1429]" box="[498,813,1160,1188]" pageId="88" pageNumber="89">
|
||
<heading bold="true" box="[498,813,1160,1188]" fontSize="11" level="3" pageId="88" pageNumber="89" reason="6">
|
||
<emphasis bold="true" box="[498,813,1160,1188]" italics="true" pageId="88" pageNumber="89">Antorbital fossa length</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1542,1160,1429]" pageId="88" pageNumber="89">
|
||
Geometric morphometric comparisons recover a shortening of the antorbital fossa during growth, a trend that is ancestral at least for
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[1103,1224,1244,1270]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="88" pageNumber="89" phylum="Chordata" rank="order">Saurischia</taxonomicName>
|
||
, aside from an apomorphic lengthening in
|
||
<emphasis box="[828,975,1284,1310]" italics="true" pageId="88" pageNumber="89">Coeolophysis</emphasis>
|
||
(
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[993,1368,1283,1310]" journalOrPublisher="PeerJ" pageId="88" pageNumber="89" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[993,1368,1283,1310]" italics="true" pageId="88" pageNumber="89">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<figureCitation box="[1383,1463,1284,1310]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="88" pageNumber="89">Fig. 33</figureCitation>
|
||
). This landmark-based comparison has not been done for
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1166,1327,1323,1349]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1166,1327,1323,1349]" italics="true" pageId="88" pageNumber="89">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1369,1437,1324,1349]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1369,1437,1324,1349]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, but it is predicted here that it will be recovered in both taxa and, hence, the trend is plesiomorphic for Archosauriformes.
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1542,1466,1934]" box="[498,743,1466,1494]" pageId="88" pageNumber="89">
|
||
<heading bold="true" box="[498,743,1466,1494]" fontSize="11" level="3" pageId="88" pageNumber="89" reason="6">
|
||
<emphasis bold="true" box="[498,743,1466,1494]" italics="true" pageId="88" pageNumber="89">Jaw joint position</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="88.[498,1542,1466,1934]" pageId="88" pageNumber="89">
|
||
Geometric morphometric comparisons have recovered the dorsoventral and rostrocaudal growth trends of the cranial jaw joint for
|
||
<taxonomicName authority="(Ezcurra & Butler, 2015)" baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[991,1452,1549,1576]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[991,1152,1549,1575]" italics="true" pageId="88" pageNumber="89">Proterosuchus</emphasis>
|
||
(
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1171,1442,1549,1576]" journalOrPublisher="Journal of Anatomy" pageId="88" pageNumber="89" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1171,1442,1549,1576]" italics="true" pageId="88" pageNumber="89">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="(Foth, Hendrick & Ezcurra, 2016)" baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[498,1012,1589,1616]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="88" pageNumber="89" phylum="Chordata" rank="order">
|
||
Saurischia (
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" box="[634,1001,1589,1616]" journalOrPublisher="PeerJ" pageId="88" pageNumber="89" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis box="[634,1001,1589,1616]" italics="true" pageId="88" pageNumber="89">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Optimization of jaw joint position recovers a caudalward shift as the ancestral state for Archosauriformes, with two independently acquired apomorphic rostral shifts in
|
||
<taxonomicName authorityName="Owen" authorityYear="1854" box="[945,1131,1669,1695]" class="Reptilia" family="Massospondylidae" genus="Massospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[945,1131,1669,1695]" italics="true" pageId="88" pageNumber="89">Massospondylus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[1192,1314,1669,1695]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1192,1314,1669,1695]" italics="true" pageId="88" pageNumber="89">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[1333,1412,1669,1695]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="88" pageNumber="89">Fig. 33</figureCitation>
|
||
). It is predicted here that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[727,795,1710,1735]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[727,795,1710,1735]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
will be found to have the same caudalward shift with
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[1427,1535,1709,1735]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1427,1535,1709,1735]" italics="true" pageId="88" pageNumber="89">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
. Likewise, landmark-based comparisons (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[977,1248,1748,1775]" journalOrPublisher="Journal of Anatomy" pageId="88" pageNumber="89" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[977,1248,1748,1775]" italics="true" pageId="88" pageNumber="89">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Foth C & Hendrick BP & Ezcurra MD" journalOrPublisher="PeerJ" pageId="88" pageNumber="89" pagination="e 1589" part="4" refId="ref53469" refString="Foth C, Hendrick BP, Ezcurra MD. 2016. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4: e 1589 DOI 10.7717 / peerj. 1589." title="Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs" type="journal article" year="2016">
|
||
<emphasis italics="true" pageId="88" pageNumber="89">Foth, Hendrick & Ezcurra, 2016</emphasis>
|
||
</bibRefCitation>
|
||
) show that a ventralward shift of the jaw joint is plesiomorphic for Archosauriformes, with an apomorphic dorsalward shift in
|
||
<taxonomicName box="[1202,1335,1828,1854]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1202,1335,1828,1854]" italics="true" pageId="88" pageNumber="89">Coelophysis</emphasis>
|
||
</taxonomicName>
|
||
and no dorsoventral shift in
|
||
<taxonomicName box="[746,945,1868,1894]" class="Reptilia" family="Megalosauridae" genus="Dubreuillosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="88" pageNumber="89" phylum="Chordata" rank="genus">
|
||
<emphasis box="[746,945,1868,1894]" italics="true" pageId="88" pageNumber="89">Dubreuillosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[963,1042,1868,1894]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="88" pageNumber="89">Fig. 33</figureCitation>
|
||
). It is predicted here that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1354,1422,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1354,1422,1869,1894]" italics="true" pageId="88" pageNumber="89">T. rex</emphasis>
|
||
</taxonomicName>
|
||
will be found to share a ventralward shift of the jaw joint with its sister species,
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[1358,1467,1908,1934]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="88" pageNumber="89" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[1358,1467,1908,1934]" italics="true" pageId="88" pageNumber="89">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1542,233,542]" box="[498,1177,233,261]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,1177,233,261]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,1177,233,261]" italics="true" pageId="89" pageNumber="90">Dorsoventral deepening of the mandibular ramus</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1542,233,542]" pageId="89" pageNumber="90">
|
||
The ontogenetic deepening of the mandibular ramus in archosaurs is evidenced by the entire ramus and by the external mandibular fenestra. In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1178,1247,317,342]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1178,1247,317,342]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and crocodylians the external mandibular fenestra and the mandibular ramus increase in dorsoventral height (
|
||
<bibRefCitation author="Monteiro LR & Soares M." box="[508,797,396,423]" journalOrPublisher="Herpetologica" pageId="89" pageNumber="90" pagination="62 - 69" part="53" refId="ref55228" refString="Monteiro LR, Soares M. 1997. Allometric analysis of the ontogenetic variation and evolution of the skull in Caiman Spix, 1825 (Crocodylia: Alligatoridae). Herpetologica 53: 62 - 69." title="Allometric analysis of the ontogenetic variation and evolution of the skull in Caiman Spix, 1825 (Crocodylia: Alligatoridae)" type="journal article" year="1997">
|
||
<emphasis box="[508,797,396,423]" italics="true" pageId="89" pageNumber="90">Monteiro & Soares, 1997</emphasis>
|
||
</bibRefCitation>
|
||
). Optimization of the trend of increasing mandible height onto the phylogenetic topology recovers it as a synapomorphy of Archosauria (
|
||
<figureCitation box="[1373,1452,436,462]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="89" pageNumber="90">Fig. 33</figureCitation>
|
||
). The lower jaw was not included in the study of
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1065,1226,476,502]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1065,1226,476,502]" italics="true" pageId="89" pageNumber="90">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
ontogeny and so it is unknown if the mandible deepened as well (
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[1023,1293,516,542]" journalOrPublisher="Journal of Anatomy" pageId="89" pageNumber="90" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[1023,1293,516,542]" italics="true" pageId="89" pageNumber="90">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,575,884]" box="[498,951,575,603]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,951,575,603]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,951,575,603]" italics="true" pageId="89" pageNumber="90">Increase in maxillary tooth count</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,575,884]" pageId="89" pageNumber="90">
|
||
An ontogenetic increase in tooth count of archosauriforms is seen in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1319,1388,620,645]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1319,1388,620,645]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="(Ezcurra & Butler, 2015)" baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[498,960,659,685]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[498,659,659,685]" italics="true" pageId="89" pageNumber="90">Proterosuchus</emphasis>
|
||
(
|
||
<bibRefCitation author="Ezcurra MD & Butler RJ" box="[679,949,659,685]" journalOrPublisher="Journal of Anatomy" pageId="89" pageNumber="90" pagination="387 - 402" part="226" refId="ref53386" refString="Ezcurra MD, Butler RJ. 2015. Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi. Journal of Anatomy 226 (5): 387 - 402 DOI 10.1111 / joa. 12300." title="Post-hatchling ontogeny in the Early Triassic diapsid reptile Proterosuchus fergusi" type="journal article" year="2015">
|
||
<emphasis box="[679,949,659,685]" italics="true" pageId="89" pageNumber="90">Ezcurra & Butler, 2015</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
; in contrast, tooth number is ontogenetically constant in extant crocodylians (
|
||
<bibRefCitation author="Brown CM & VanBuren CS & Larson DW & Brink KS & Campione NE & Vavrek MJ & Evans DC" box="[885,1098,699,725]" journalOrPublisher="Journal of Anatomy" pageId="89" pageNumber="90" pagination="322 - 333" part="226" refId="ref52355" refString="Brown CM, VanBuren CS, Larson DW, Brink KS, Campione NE, Vavrek MJ, Evans DC. 2015. Tooth counts through growth in diapsid reptiles: implications for interpreting individual and sizerelated variation in the fossil record. Journal of Anatomy 226 (4): 322 - 333 DOI 10.1111 / joa. 12280." title="Tooth counts through growth in diapsid reptiles: implications for interpreting individual and sizerelated variation in the fossil record" type="journal article" year="2015">
|
||
<emphasis box="[885,1098,699,725]" italics="true" pageId="89" pageNumber="90">Brown et al., 2015</emphasis>
|
||
</bibRefCitation>
|
||
). Optimization of the trend in tooth count onto the phylogenetic topology recovers an increase in number as plesiomorphic for Archosauriformes (
|
||
<figureCitation box="[770,849,778,804]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="89" pageNumber="90">Fig. 33</figureCitation>
|
||
).The presence of ontogenetic tooth increase in other dinosaurs and the extant sister clade of Archosauria, Squamata, indicates that the pattern seen in crocodylians is the apomorphic condition (
|
||
<bibRefCitation author="Brown CM & VanBuren CS & Larson DW & Brink KS & Campione NE & Vavrek MJ & Evans DC" box="[1100,1312,858,884]" journalOrPublisher="Journal of Anatomy" pageId="89" pageNumber="90" pagination="322 - 333" part="226" refId="ref52355" refString="Brown CM, VanBuren CS, Larson DW, Brink KS, Campione NE, Vavrek MJ, Evans DC. 2015. Tooth counts through growth in diapsid reptiles: implications for interpreting individual and sizerelated variation in the fossil record. Journal of Anatomy 226 (4): 322 - 333 DOI 10.1111 / joa. 12280." title="Tooth counts through growth in diapsid reptiles: implications for interpreting individual and sizerelated variation in the fossil record" type="journal article" year="2015">
|
||
<emphasis box="[1100,1312,858,884]" italics="true" pageId="89" pageNumber="90">Brown et al., 2015</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,917,1187]" box="[498,631,917,945]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,631,917,945]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,631,917,945]" italics="true" pageId="89" pageNumber="90">Bite force</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,917,1187]" pageId="89" pageNumber="90">
|
||
A growth-related increase in bite forces is seen in crocodylians and in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1304,1370,962,987]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1304,1370,962,987]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, a trend that is continued into adulthood (
|
||
<bibRefCitation author="Bates KT & Falkingham PL" box="[818,1127,1001,1027]" journalOrPublisher="Biology Letters" pageId="89" pageNumber="90" pagination="660 - 664" part="8" refId="ref52110" refString="Bates KT, Falkingham PL. 2012. Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Biology Letters 8 (4): 660 - 664 DOI 10.1098 / rsbl. 2012.0056." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">
|
||
<emphasis box="[818,1127,1001,1027]" italics="true" pageId="89" pageNumber="90">Bates & Falkingham, 2012</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Erickson GM & Lappin AK & Vliet KA" box="[1142,1502,1001,1027]" journalOrPublisher="Journal of Zoology" pageId="89" pageNumber="90" pagination="317 - 327" part="260" refId="ref53166" refString="Erickson GM, Lappin AK, Vliet KA. 2003. The ontogeny of bite-force performance in American Alligator (Alligator mississippiensis). Journal of Zoology 260 (3): 317 - 327 DOI 10.1017 / S 0952836903003819." title="The ontogeny of bite-force performance in American Alligator (Alligator mississippiensis)" type="journal article" year="2003">
|
||
<emphasis box="[1142,1502,1001,1027]" italics="true" pageId="89" pageNumber="90">Erickson, Lappin & Vliet, 2003</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Erickson GM & Gignac PM & Lappin AK & Vliet KA & Brueggen JD & Webb GJW" box="[498,733,1041,1067]" journalOrPublisher="Journal of Zoology" pageId="89" pageNumber="90" pagination="48 - 55" part="292" refId="ref53338" refString="Erickson GM, Gignac PM, Lappin AK, Vliet KA, Brueggen JD, Webb GJW. 2013. A comparative analysis of ontogenetic bite-force scaling among Crocodylia. Journal of Zoology 292 (1): 48 - 55 DOI 10.1111 / jzo. 12081." title="A comparative analysis of ontogenetic bite-force scaling among Crocodylia" type="journal article" year="2013">
|
||
<emphasis box="[498,733,1041,1067]" italics="true" pageId="89" pageNumber="90">Erickson et al., 2013</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[748,1037,1041,1067]" journalOrPublisher="Journal of Zoology" pageId="89" pageNumber="90" pagination="132 - 142" part="295" refId="ref53694" refString="Gignac PM, Erickson GM. 2014. Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators. Journal of Zoology 295 (2): 132 - 142 DOI 10.1111 / jzo. 12187." title="Ontogenetic changes in dental form and tooth pressures facilitate developmental niche shifts in American alligators" type="journal article" year="2014">
|
||
<emphasis box="[748,1037,1041,1067]" italics="true" pageId="89" pageNumber="90">Gignac & Erickson, 2014</emphasis>
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Gignac PM & Erickson GM" box="[1052,1107,1041,1067]" journalOrPublisher="Scientific Reports" pageId="89" pageNumber="90" pagination="479" part="7" refId="ref53777" refString="Gignac PM, Erickson GM. 2017. The biomechanics behind extreme osteophagy in Tyrannosaurus rex. Scientific Reports 7 (1): 479 DOI 10.1038 / s 41598 - 017 - 02161 - w." title="The biomechanics behind extreme osteophagy in Tyrannosaurus rex" type="journal article" year="2017">
|
||
<emphasis box="[1052,1107,1041,1067]" italics="true" pageId="89" pageNumber="90">2017</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<emphasis box="[1121,1401,1041,1067]" italics="true" pageId="89" pageNumber="90">Gignac & O’ Brien, 2016</emphasis>
|
||
). Optimization of this trend onto the phylogenetic topology recovers increasing bite force as a synapomorphy of Archosauria; it is expected that this trend is pleiomorphic for Archosauriformes, once data are obtained for
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[1043,1204,1160,1186]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1043,1204,1160,1186]" italics="true" pageId="89" pageNumber="90">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
or other basal taxa.
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1220,1449]" box="[498,828,1220,1248]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,828,1220,1248]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,828,1220,1248]" italics="true" pageId="89" pageNumber="90">Cephalic ornamentation</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1220,1449]" pageId="89" pageNumber="90">
|
||
Growth-related reduction in cephalic ornamentation is seen in
|
||
<taxonomicName authority="(Horner & Goodwin, 2006)" baseAuthorityName="Horner & Goodwin" baseAuthorityYear="2006" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1245,1375,1264,1289]" italics="true" pageId="89" pageNumber="90">Triceratops</emphasis>
|
||
(
|
||
<bibRefCitation author="Horner JR & Goodwin MB" journalOrPublisher="Proceedings of the Royal Society of London: Biology" pageId="89" pageNumber="90" pagination="2757 - 2761" part="273" refId="ref54486" refString="Horner JR, Goodwin MB. 2006. Major cranial changes during Triceratops ontogeny. Proceedings of the Royal Society of London: Biology 273 (1602): 2757 - 2761 DOI 10.1098 / rspb. 2006.3643." title="Major cranial changes during Triceratops ontogeny" type="journal article" year="2006">
|
||
<emphasis italics="true" pageId="89" pageNumber="90">Horner & Goodwin, 2006</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="(Horner & Goodwin, 2009)" baseAuthorityName="Horner & Goodwin" baseAuthorityYear="2009" box="[698,1257,1303,1330]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[698,929,1303,1329]" italics="true" pageId="89" pageNumber="90">Pachycephalosaurus</emphasis>
|
||
(
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[948,1246,1303,1330]" journalOrPublisher="PLOS ONE" pageId="89" pageNumber="90" pagination="e 7626" part="4" refId="ref54525" refString="Horner JR, Goodwin MB. 2009. Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLOS ONE 4 (10): e 7626 DOI 10.1371 / journal. pone. 0007626." title="Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus" type="journal article" year="2009">
|
||
<emphasis box="[948,1246,1303,1330]" italics="true" pageId="89" pageNumber="90">Horner & Goodwin, 2009</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, and in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1356,1425,1304,1329]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1356,1425,1304,1329]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Optimization of the growth trend onto the phylogenetic topology recovers this pattern as synapomorphic for
|
||
<taxonomicName authorityName="Owen" authorityYear="1842" box="[731,860,1383,1409]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
(
|
||
<figureCitation box="[879,959,1383,1409]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="89" pageNumber="90">Fig. 33</figureCitation>
|
||
); it is predicted that this trend will be an archosauriform synapomorphy as appropriate data are documented.
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1482,1752]" box="[498,808,1482,1510]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,808,1482,1510]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,808,1482,1510]" italics="true" pageId="89" pageNumber="90">Appendicular skeleton</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1482,1752]" pageId="89" pageNumber="90">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[498,567,1527,1552]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[498,567,1527,1552]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
shares two postcranial growth changes with
|
||
<taxonomicName authority="(Griffin, 2018)" baseAuthorityName="Griffin" baseAuthorityYear="2018" box="[1099,1409,1526,1552]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1099,1232,1526,1552]" italics="true" pageId="89" pageNumber="90">Coelophysis</emphasis>
|
||
(
|
||
<bibRefCitation author="Griffin CT" box="[1251,1398,1526,1552]" journalOrPublisher="Journal of Anatomy" pageId="89" pageNumber="90" pagination="604 - 640" part="232" refId="ref53894" refString="Griffin CT. 2018. Developmental patterns and variation among early theropods. Journal of Anatomy 232 (4): 604 - 640 DOI 10.1111 / joa. 12775." title="Developmental patterns and variation among early theropods" type="journal article" year="2018">
|
||
<emphasis box="[1251,1398,1526,1552]" italics="true" pageId="89" pageNumber="90">Griffin, 2018</emphasis>
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, which indicates synapomorphic growth changes that possibly unite Neotheropoda. These include the thickened deltopectoral crest of the humerus and enhancement of the trochanteric shelf of the femur. When optimized onto the phylogenetic topology, both trends are recovered as synapomorphies of Neotheropoda; it is predicted that these trends will be found to be synapomorphies of Archosauriformes.
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1784,1934]" box="[498,631,1784,1812]" pageId="89" pageNumber="90">
|
||
<heading bold="true" box="[498,631,1784,1812]" fontSize="11" level="3" pageId="89" pageNumber="90" reason="6">
|
||
<emphasis bold="true" box="[498,631,1784,1812]" italics="true" pageId="89" pageNumber="90">Summary</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="89.[498,1541,1784,1934]" lastBlockId="90.[498,1542,234,659]" lastPageId="90" lastPageNumber="91" pageId="89" pageNumber="90">
|
||
Taken together, the ancestral growth pattern of the archosauriform skull that was inherited by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[535,604,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="89" pageNumber="90" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[535,604,1869,1894]" italics="true" pageId="89" pageNumber="90">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, was marked by (1) increase in height of the entire skull, (2) rostrocaudal and dorsoventral expansion of the adductor chamber, (3) increase in skull width, (4) caudalward shift of the jaw joint, (5) ventralward shift of the jaw joint, and an (6) increase in maxillary tooth number (
|
||
<figureCitation box="[933,1012,274,300]" captionStart="Figure 33" captionStartId="87.[524,589,1254,1276]" captionTargetBox="[498,1542,234,1242]" captionTargetId="graphics-114@87.[498,1542,261,1241]" captionTargetPageId="87" captionText="Figure 33 A simplified cladogram of living and extinct Archosauriformes showing 13 cranial and postcranial growth changes that are optimized as synapomorphies. Most of the growth changes are ancestral for Archosauriformes. The position of several characters at progressively exclusive clades is almost certainly an artifact of missing data (e.g., increase in mandible height, enlargement of muscle attachments, etc.) and they are predicted to be synapomorphic for Archosauriformes once the appropriate data are acquired. This comparison shows that highly derived species such as Tyrannosaurus rex do not deviate from the ancestral growth trends that first evolved in significantly smaller taxa. See text for sources; see Table 23 for the distribution of character states among the taxa. Full-size DOI: 10.7717/peerj.9192/fig-33" figureDoi="http://doi.org/10.5281/zenodo.5825040" httpUri="https://zenodo.org/record/5825040/files/figure.png" pageId="90" pageNumber="91">Fig. 33</figureCitation>
|
||
). The ancestral growth pattern of the archosaurian skull that was inherited by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[979,1048,315,340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[979,1048,315,340]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
included (1) increase in mandible height and (2) an increase in bite force. The ancestral growth pattern of the dinosaurian skull that was inherited directly by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[787,853,394,419]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[787,853,394,419]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was the reduction in cephalic ornamentation. The ancestral growth pattern of the saurischian skull that was inherited by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1193,1259,434,459]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1193,1259,434,459]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
included the decrease in the length of the antorbital fossa. The ancestral growth pattern of Neotheropoda that was inherited directly by
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[787,853,514,539]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[787,853,514,539]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
included increase in size of (1) the deltopectoral crest of the humerus and (2) the trochanteric crest of the femur. It is expected that the more exclusive trends will eventually be seen in
|
||
<taxonomicName baseAuthorityName="Ezcurra & Butler" baseAuthorityYear="2015" box="[885,1046,593,619]" class="Reptilia" family="Proterosuchidae" genus="Proterosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="90" pageNumber="91" phylum="Chordata" rank="genus">
|
||
<emphasis box="[885,1046,593,619]" italics="true" pageId="90" pageNumber="91">Proterosuchus</emphasis>
|
||
</taxonomicName>
|
||
and other basal archosauriformians once adequate data are obtained.
|
||
</paragraph>
|
||
<paragraph blockId="90.[498,1542,704,1413]" box="[498,974,704,734]" pageId="90" pageNumber="91">
|
||
<heading bold="true" box="[498,974,704,734]" fontSize="12" level="2" pageId="90" pageNumber="91" reason="0">
|
||
<emphasis bold="true" box="[498,974,704,734]" pageId="90" pageNumber="91">Early onset of adult skull shape</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="90.[498,1542,704,1413]" pageId="90" pageNumber="91">
|
||
The unique skull shape of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[813,882,750,775]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[813,882,750,775]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, in having a mediolaterally narrow snout but wide orbitotemporal region (
|
||
<figureCitation box="[776,866,789,816]" captionStart="Figure 2" captionStartId="13.[524,589,1473,1495]" captionTargetBox="[475,1542,220,1456]" captionTargetId="graphics-14@13.[498,1541,247,1461]" captionTargetPageId="13" captionText="Figure 2 Ontogram of Tyrannosaurus rex showing growth stages, synontomorphies, individual variation, individual specimens, and chronological ages. Arrowhead points to the most mature spe- cimen and the direction of the entire ontogenetic axis; that is, the least mature specimen is at the lower left whereas the most mature specimen is at the upper right. Asterisk indicates the type specimen. Individual variation occurs as progressions until young adulthood, where reversals are first seen. The maximum amount of change occurs at growth stages 5 and 6, which corresponds to the transition from a long and low skull and jaws to a deep and stout skull frame; this event, marked by the concentration of an extreme number of changes, is evidence that the ontogeny of T. rex is metamorphic (sensu Rose & Reiss, 1993). Each circle represents a numbered growth stage; these numbers do not correspond to those seen in Fig. 12. The star at growth stage 7 marks the ~3,000 kg threshold that separates T. rex from its closest, but smaller, relatives. Color key: red, small juveniles; orange, large juveniles; yellow, subadults; green, young adults; blue, adults; violet, senescent adults. See text for definition of growth categories. Skulls are to scale; AMNH FARB 5027 is scaled to a premaxilla to quadrate length of 1.3 m. Full-size DOI: 10.7717/peerj.9192/fig-2" figureDoi="http://doi.org/10.5281/zenodo.5824976" httpUri="https://zenodo.org/record/5824976/files/figure.png" pageId="90" pageNumber="91">Figs. 12</figureCitation>
|
||
and
|
||
<figureCitation box="[927,955,789,815]" captionStart="Figure 26" captionStartId="68.[524,589,904,926]" captionTargetBox="[498,1541,234,890]" captionTargetPageId="68" captionText="Figure 26 Heat maps of the ontogenetic changes seen in the skull and mandible of Tyrannosaurus rex. Illustrations show per centage of the total number of unambiguously optimized synontomorphies per bone (A) and functional module (B). Darker shades of gray indicate higher proportions of growth change, whereas lighter shades indicate lower proportions of change. The results show that the greatest amount of growth changes are at the lacrimal (A) or along the dorsal skull roof (B). Hatchure indicates empty space; stipple indicates unprepared matrix. Full-size DOI: 10.7717/peerj.9192/fig-26" figureDoi="http://doi.org/10.5281/zenodo.5825026" httpUri="https://zenodo.org/record/5825026/files/figure.png" pageId="90" pageNumber="91">26</figureCitation>
|
||
), is seen in the youngest of the most complete specimens in the sample (e.g.,
|
||
<materialsCitation box="[859,1014,829,856]" collectionCode="CMNH" pageId="90" pageNumber="91" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
) that is approximately 40% of adult skull length. This phenomenon of the presence of the adult shape in juveniles (i.e., developmental capture; sensu
|
||
<bibRefCitation author="Padian K." box="[911,1064,909,935]" editor="Thomason JJ" journalOrPublisher="Cambridge: University of Cambridge Press" pageId="90" pageNumber="91" pagination="264 - 277" refId="ref55474" refString="Padian K. 1995. Form versus function: the evolution of a dialectic. In: Thomason JJ, ed. Functional Morphology in Vertebrate Paleontology. Cambridge: University of Cambridge Press, 264 - 277." title="Form versus function: the evolution of a dialectic" type="book chapter" volumeTitle="Functional Morphology in Vertebrate Paleontology" year="1995">
|
||
<emphasis box="[911,1064,909,935]" italics="true" pageId="90" pageNumber="91">Padian, 1995</emphasis>
|
||
</bibRefCitation>
|
||
) is not unusual among archosaurs; for example, it is also seen in caimans (
|
||
<bibRefCitation author="Monteiro LR & Soares M." box="[906,1189,949,975]" journalOrPublisher="Herpetologica" pageId="90" pageNumber="91" pagination="62 - 69" part="53" refId="ref55228" refString="Monteiro LR, Soares M. 1997. Allometric analysis of the ontogenetic variation and evolution of the skull in Caiman Spix, 1825 (Crocodylia: Alligatoridae). Herpetologica 53: 62 - 69." title="Allometric analysis of the ontogenetic variation and evolution of the skull in Caiman Spix, 1825 (Crocodylia: Alligatoridae)" type="journal article" year="1997">
|
||
<emphasis box="[906,1189,949,975]" italics="true" pageId="90" pageNumber="91">Monteiro & Soares, 1997</emphasis>
|
||
</bibRefCitation>
|
||
). This phenomenon is notable in that the adult skull shape in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[869,938,990,1015]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[869,938,990,1015]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is derived in contrast to its closest relatives, but it ontogenetically precedes the occurrence of the suite of other evolutionary novelties that sets
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[549,618,1069,1094]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[549,618,1069,1094]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
apart from
|
||
<taxonomicName baseAuthorityName="Foth, Hendrick & Ezcurra" baseAuthorityYear="2016" box="[762,871,1068,1094]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis box="[762,871,1068,1094]" italics="true" pageId="90" pageNumber="91">T. bataar</emphasis>
|
||
</taxonomicName>
|
||
and other advanced tyrannosaurines.
|
||
</paragraph>
|
||
<paragraph blockId="90.[498,1542,704,1413]" pageId="90" pageNumber="91">
|
||
This observation is evidence that skull shape in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1085,1152,1109,1134]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1085,1152,1109,1134]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is not recapitulatorily locked to a later growth stage; rather, continuity is seen in skull function and behavior between juvenile and adult
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[719,787,1189,1214]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[719,787,1189,1214]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and niche partitioning between those small and large growth stages was primarily governed by increased bite force instead of fundamental differences in sensorimotor integration and functional morphology (aside from the consequences of the corresponding increase in skull and jaw height) and mode of forage. From its earliest growth stage,
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[663,732,1348,1373]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[663,732,1348,1373]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
had already fundamentally departed from its ancestral skull morphology and, presumably, function.
|
||
</paragraph>
|
||
<paragraph blockId="90.[498,1542,1458,1889]" box="[498,1193,1458,1489]" pageId="90" pageNumber="91">
|
||
<heading bold="true" box="[498,1193,1458,1489]" fontSize="12" level="2" pageId="90" pageNumber="91" reason="0">
|
||
<emphasis bold="true" box="[498,1193,1458,1489]" pageId="90" pageNumber="91">Juvenile to ‘Adult’ skull morphotype transition</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="90.[498,1542,1458,1889]" lastBlockId="91.[498,1530,899,966]" lastPageId="91" lastPageNumber="92" pageId="90" pageNumber="91">
|
||
The results found here show that the extreme ontogenetic transition in skull shape of
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1476,1542,1505,1530]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1476,1542,1505,1530]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is often mischaracterized as between juvenile to adult, whereas the switch is actually much earlier, between juvenile and subadult. In living crocodiles, transitions in skull shape mark changes in diet (e.g.,
|
||
<taxonomicName baseAuthorityName="Cuvier" baseAuthorityYear="1807" box="[888,996,1624,1650]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="acutus">
|
||
<emphasis box="[888,996,1624,1650]" italics="true" pageId="90" pageNumber="91">C. acutus</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<bibRefCitation author="Platt SG & Rainwater TR & Thorbjarnarson JB & Martin D." box="[1010,1198,1624,1650]" journalOrPublisher="Salamandra" pageId="90" pageNumber="91" pagination="179 - 192" part="47" refId="ref55690" refString="Platt SG, Rainwater TR, Thorbjarnarson JB, Martin D. 2011. Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize. Salamandra 47: 179 - 192." title="Size estimation, morphometrics, sex ratio, sexual size dimorphism, and biomass of Crocodylus acutus in the coastal zone of Belize" type="journal article" year="2011">
|
||
<emphasis box="[1010,1198,1624,1650]" italics="true" pageId="90" pageNumber="91">Platt et al., 2011</emphasis>
|
||
</bibRefCitation>
|
||
), and the same is doubtlessly true in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[586,655,1664,1689]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[586,655,1664,1689]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Aside from skull shape in living crocodiles (e.g.,
|
||
<taxonomicName box="[1244,1375,1664,1690]" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="niloticus">
|
||
<emphasis box="[1244,1375,1664,1690]" italics="true" pageId="90" pageNumber="91">C. niloticus</emphasis>
|
||
</taxonomicName>
|
||
), size is an important factor that drives juveniles from their natal river range to a lacustrine habitat (
|
||
<bibRefCitation author="Hutton J." box="[508,662,1743,1769]" journalOrPublisher="American Zoologist" pageId="90" pageNumber="91" pagination="1033 - 1049" part="29" refId="ref54732" refString="Hutton J. 1989. Movements, home range, dispersal and the separation of size classes in Nile crocodiles. American Zoologist 29 (3): 1033 - 1049 DOI 10.1093 / icb / 29.3.1033." title="Movements, home range, dispersal and the separation of size classes in Nile crocodiles" type="journal article" year="1989">
|
||
<emphasis box="[508,662,1743,1769]" italics="true" pageId="90" pageNumber="91">Hutton, 1989</emphasis>
|
||
</bibRefCitation>
|
||
); likewise, the extreme transition in skull shape in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1255,1323,1744,1769]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="90" pageNumber="91" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1255,1323,1744,1769]" italics="true" pageId="90" pageNumber="91">T. rex</emphasis>
|
||
</taxonomicName>
|
||
might also mark a dispersal stage with movement from, say, a closed habitat that provided protection from adults to an open setting where safety from conspecifics was no longer a priority. Presumably, the deep-skulled morphotype resulted in an expansion of range among subadults and adults, where females would be limited to nesting areas and males would have been more widely distributed across the landscape (cf.
|
||
<bibRefCitation author="Hutton J." box="[1208,1362,939,965]" journalOrPublisher="American Zoologist" pageId="91" pageNumber="92" pagination="1033 - 1049" part="29" refId="ref54732" refString="Hutton J. 1989. Movements, home range, dispersal and the separation of size classes in Nile crocodiles. American Zoologist 29 (3): 1033 - 1049 DOI 10.1093 / icb / 29.3.1033." title="Movements, home range, dispersal and the separation of size classes in Nile crocodiles" type="journal article" year="1989">
|
||
<emphasis box="[1208,1362,939,965]" italics="true" pageId="91" pageNumber="92">Hutton, 1989</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-Table-UUID="DF5C84E2FF9FFFF3FDEBFF70327DFE97" httpUri="http://table.plazi.org/id/DF5C84E2FF9FFFF3FDEBFF70327DFE97" pageId="91" pageNumber="92" startId="91.[525,580,248,269]" targetBox="[516,1499,320,846]" targetIsTable="true" targetPageId="91">
|
||
<paragraph blockId="91.[525,1515,248,300]" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[525,1515,248,270]" pageId="91" pageNumber="92">
|
||
Table 24 Predicted LAG counts for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[886,1072,249,270]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="91" pageNumber="92" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis bold="true" box="[886,1072,249,270]" italics="true" pageId="91" pageNumber="92">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
specimens extrapolated from the ontogram.
|
||
</emphasis>
|
||
Estimated ages are indicated by a tilde (~), histologically aged specimens are in boldface.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="91" pageNumber="92">
|
||
<table box="[516,1499,320,846]" gridcols="2" gridrows="12" pageId="91" pageNumber="92">
|
||
<tr box="[516,1499,320,371]" gridrow="0" pageId="91" pageNumber="92">
|
||
<th box="[516,1404,320,371]" gridcol="0" gridrow="0" pageId="91" pageNumber="92">Specimen</th>
|
||
<th box="[1428,1499,320,371]" gridcol="1" gridrow="0" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1467,320,342]" pageId="91" pageNumber="92">Age</emphasis>
|
||
<emphasis bold="true" box="[1428,1494,349,371]" pageId="91" pageNumber="92">(years)</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr box="[516,1499,390,412]" gridrow="1" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,390,412]" gridcol="0" gridrow="1" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,650,390,412]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,390,412]" gridcol="1" gridrow="1" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1440,390,411]" pageId="91" pageNumber="92">2</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,428,450]" gridrow="2" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,428,450]" gridcol="0" gridrow="2" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,709,428,450]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47787" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5050">AMNH FARB 5050</materialsCitation>
|
||
,
|
||
<materialsCitation box="[719,846,428,450]" collectionCode="CMNH" pageId="91" pageNumber="92" specimenCode="CMNH 7541">CMNH 7541</materialsCitation>
|
||
,
|
||
<materialsCitation box="[857,972,428,449]" collectionCode="DDM" pageId="91" pageNumber="92" specimenCode="DDM 344.1">DDM 344.1</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,428,450]" gridcol="1" gridrow="2" pageId="91" pageNumber="92">
|
||
~3
|
||
<emphasis box="[1453,1465,428,450]" italics="true" pageId="91" pageNumber="92">–</emphasis>
|
||
12
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,465,487]" gridrow="3" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,465,487]" gridcol="0" gridrow="3" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,673,465,487]" collectionCode="BMRP" pageId="91" pageNumber="92" specimenCode="BMRP 2002.4.1">BMRP 2002.4.1</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,465,487]" gridcol="1" gridrow="3" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1452,465,486]" pageId="91" pageNumber="92">13</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,503,524]" gridrow="4" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,503,524]" gridcol="0" gridrow="4" pageId="91" pageNumber="92">BMRP 2006.6.4</td>
|
||
<td box="[1428,1499,503,524]" gridcol="1" gridrow="4" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1452,503,524]" pageId="91" pageNumber="92">15</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,540,563]" gridrow="5" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,540,563]" gridcol="0" gridrow="5" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,634,541,562]" collectionCode="RSM" pageId="91" pageNumber="92" specimenCode="RSM 2990.1">RSM 2990.1</materialsCitation>
|
||
,
|
||
<materialsCitation box="[645,775,540,562]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,540,563]" gridcol="1" gridrow="5" pageId="91" pageNumber="92">
|
||
~15
|
||
<emphasis box="[1464,1476,541,563]" italics="true" pageId="91" pageNumber="92">–</emphasis>
|
||
17
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,578,600]" gridrow="6" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,578,600]" gridcol="0" gridrow="6" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,627,578,600]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
,
|
||
<materialsCitation box="[639,840,578,599]" collectionCode="TMP" pageId="91" pageNumber="92" specimenCode="TMP 1981.006.0001">TMP 1981.006.0001</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,578,600]" gridcol="1" gridrow="6" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1452,578,599]" pageId="91" pageNumber="92">18</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,616,638]" gridrow="7" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,616,638]" gridcol="0" gridrow="7" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,709,616,638]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
,
|
||
<materialsCitation box="[719,861,616,638]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,616,638]" gridcol="1" gridrow="7" pageId="91" pageNumber="92">
|
||
~18
|
||
<emphasis box="[1464,1476,616,638]" italics="true" pageId="91" pageNumber="92">–</emphasis>
|
||
22
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,653,675]" gridrow="8" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,653,675]" gridcol="0" gridrow="8" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,717,653,675]" collectionCode="TMP" pageId="91" pageNumber="92" specimenCode="TMP 1981.006.0001">TMP 1981.012.0001</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,653,675]" gridcol="1" gridrow="8" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1452,653,674]" pageId="91" pageNumber="92">22</emphasis>
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,691,713]" gridrow="9" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,691,713]" gridcol="0" gridrow="9" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,656,691,713]" collectionCode="UWBM" pageId="91" pageNumber="92" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,691,713]" gridcol="1" gridrow="9" pageId="91" pageNumber="92">~22</td>
|
||
</tr>
|
||
<tr box="[516,1499,729,809]" gridrow="10" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,729,809]" gridcol="0" gridrow="10" pageId="91" pageNumber="92">
|
||
<materialsCitation box="[516,709,729,751]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[719,912,729,751]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47763" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5029">AMNH FARB 5029</materialsCitation>
|
||
,
|
||
<materialsCitation box="[923,1083,729,750]" collectionCode="NHMUK" pageId="91" pageNumber="92" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1093,1185,729,750]" collectionCode="CM" pageId="91" pageNumber="92" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1195,1292,729,750]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1303,1400,729,751]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
,
|
||
<materialsCitation box="[532,628,758,779]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<materialsCitation box="[639,747,758,779]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
,
|
||
<materialsCitation box="[758,867,758,779]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
,
|
||
<materialsCitation box="[877,1009,758,779]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1020,1192,758,780]" collectionCode="NMMNH" pageId="91" pageNumber="92" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1203,1321,758,780]" collectionCode="RSM" pageId="91" pageNumber="92" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="SDSM" pageId="91" pageNumber="92" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
|
||
,
|
||
<materialsCitation box="[601,747,787,808]" collectionCode="UCMP" pageId="91" pageNumber="92" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
,
|
||
<materialsCitation box="[757,898,787,809]" collectionCode="UMNH" pageId="91" pageNumber="92" specimenCode="UMNH 11000">UMNH 11000</materialsCitation>
|
||
,
|
||
<materialsCitation box="[909,1028,787,808]" collectionCode="UWGM" pageId="91" pageNumber="92" specimenCode="UWGM 181">UWGM 181</materialsCitation>
|
||
</td>
|
||
<td box="[1428,1499,729,809]" gridcol="1" gridrow="10" pageId="91" pageNumber="92">
|
||
~23
|
||
<emphasis box="[1464,1476,729,751]" italics="true" pageId="91" pageNumber="92">–</emphasis>
|
||
27
|
||
</td>
|
||
</tr>
|
||
<tr box="[516,1499,825,846]" gridrow="11" pageId="91" pageNumber="92">
|
||
<td box="[516,1404,825,846]" gridcol="0" gridrow="11" pageId="91" pageNumber="92">FMNH PR2981</td>
|
||
<td box="[1428,1499,825,846]" gridcol="1" gridrow="11" pageId="91" pageNumber="92">
|
||
<emphasis bold="true" box="[1428,1452,825,846]" pageId="91" pageNumber="92">28</emphasis>
|
||
</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,1542,1004,1435]" box="[498,1004,1004,1034]" pageId="91" pageNumber="92">
|
||
<heading bold="true" box="[498,1004,1004,1034]" fontSize="12" level="2" pageId="91" pageNumber="92" reason="0">
|
||
<emphasis bold="true" box="[498,1004,1004,1034]" pageId="91" pageNumber="92">Ontogeny and module integration</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,1542,1004,1435]" pageId="91" pageNumber="92">
|
||
Uniform changes across the skull implies functional integration of the skull in contrast to modularity (
|
||
<bibRefCitation author="Monteiro LR & Cavalcanti MJ & Sommer HJS III." box="[646,1098,1089,1116]" journalOrPublisher="Journal of Morphology" pageId="91" pageNumber="92" pagination="53 - 62" part="231" refId="ref55265" refString="Monteiro LR, Cavalcanti MJ, Sommer HJS III. 1997. Comparative ontogenetic shape changes in the skull of Caiman species (Crocodylia, Alligatoridae). Journal of Morphology 231: 53 - 62 DOI 10.1002 / (SICI) 1097 - 4687 (199701) 231: 1 <53 :: AID-JMOR 5> 3.0. CO; 2 - P." title="Comparative ontogenetic shape changes in the skull of Caiman species (Crocodylia, Alligatoridae)" type="journal article" year="1997">
|
||
<emphasis box="[646,1098,1089,1116]" italics="true" pageId="91" pageNumber="92">Monteiro, Cavalcanti & Sommer, 1997</emphasis>
|
||
</bibRefCitation>
|
||
). In
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1157,1226,1090,1115]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="91" pageNumber="92" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1157,1226,1090,1115]" italics="true" pageId="91" pageNumber="92">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, the skull modules follow the same general pattern of change, starting with few changes (<10), followed by a peak (>10) from the 5th to 6th growth stages, followed thereafter by a low rate of change (<10) (
|
||
<tableCitation box="[672,759,1209,1235]" captionStart="Table 4" captionStartId="22.[117,172,963,984]" captionTargetBox="[108,1525,1064,1353]" captionText="Table 4 Summary of synontomorphies and individual variation in Tyrannosaurus rex organized by growth category and functional modules of the skull and jaws. Summary of the number of unambiguously optimized changes recovered for the craniomandibular skeleton of Tyrannosaurus rex, organized by functional modules (sensu Werneburg et al., 2019). Individual variation is shown in parentheses." httpUri="http://table.plazi.org/id/DF5C84E2FFD2FFBEFF93FC4B33DCFBAF" pageId="91" pageNumber="92" tableUuid="DF5C84E2FFD2FFBEFF93FC4B33DCFBAF">Table 4</tableCitation>
|
||
;
|
||
<figureCitation box="[774,854,1209,1235]" captionStart="Figure 11" captionStartId="22.[524,589,695,717]" captionTargetBox="[502,1535,236,665]" captionTargetId="figure-171@22.[497,1130,233,669]" captionTargetPageId="22" captionText="Figure 11 The frequency distribution of changes to the craniomandibular functional modules (sensu Werneburg et al., 2019) in the growth series of Tyrannosaurus rex. Growth stages are along the x-axis (corresponding to the numbered nodes of the ontogram in Fig. 2) and the y-axis corresponds to the number of synontomorphies. The onset of the changes to the skull roof, snout, mandibular ramus, and suspensorium modules occur early in growth, whereas the onset of changes to the parietal and braincase occur in adulthood. Changes continue throughout growth in all domains, aside from those to the parietal that cease at growth stage 14. Full-size DOI: 10.7717/peerj.9192/fig-11" figureDoi="http://doi.org/10.5281/zenodo.5824994" httpUri="https://zenodo.org/record/5824994/files/figure.png" pageId="91" pageNumber="92">Fig. 11</figureCitation>
|
||
). The skull roof has a peak of change at growth stage 21, showing its independence from the other modules. Beyond that instance, the modules do not tend to change independently from each other from young adulthood onwards, indicating that the modules are tightly integrated with each other, which is consistent with the akinetic hypothesis (
|
||
<bibRefCitation author="Cost IN & Middleton KM & Sellers KC & Echols MS & Witmer LM & Davis JL & Holliday CM" box="[786,973,1368,1395]" journalOrPublisher="Anatomical Record" pageId="91" pageNumber="92" pagination="1 - 19" part="303" refId="ref52906" refString="Cost IN, Middleton KM, Sellers KC, Echols MS, Witmer LM, Davis JL, Holliday CM. 2019. Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex. Anatomical Record 303 (4): 1 - 19 DOI 10.1002 / ar. 24219." title="Palatal biomechanics and its significance for cranial kinesis in Tyrannosaurus rex" type="journal article" year="2019">
|
||
<emphasis box="[786,973,1368,1395]" italics="true" pageId="91" pageNumber="92">Cost et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
) in contrast to the flexible skull hypothesis (cf.
|
||
<bibRefCitation author="Werneburg I & Esteve-Altava D & Bruno J & Ladeira MT & Diogo Rui" box="[498,764,1408,1434]" journalOrPublisher="Scientific Reports" pageId="91" pageNumber="92" pagination="1520" part="9" refId="ref56443" refString="Werneburg I, Esteve-Altava D, Bruno J, Ladeira MT, Diogo Rui. 2019. Unique skull network complexity of Tyrannosaurus rex among land vertebrate. Scientific Reports 9 (1): 1520 DOI 10.1038 / s 41598 - 018 - 37976 - 8." title="Unique skull network complexity of Tyrannosaurus rex among land vertebrate" type="journal article" year="2019">
|
||
<emphasis box="[498,764,1408,1434]" italics="true" pageId="91" pageNumber="92">Werneburg et al., 2019</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,670,1473,1503]" box="[498,670,1473,1503]" pageId="91" pageNumber="92">
|
||
<heading bold="true" box="[498,670,1473,1503]" fontSize="12" level="2" pageId="91" pageNumber="92" reason="0">
|
||
<emphasis bold="true" box="[498,670,1473,1503]" pageId="91" pageNumber="92">Predictions</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,1542,1539,1804]" pageId="91" pageNumber="92">
|
||
1. EFS will be found in
|
||
<materialsCitation box="[777,1013,1539,1566]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47761" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5027">AMNH FARB 5027</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1026,1261,1539,1566]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47763" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5029">AMNH FARB 5029</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1274,1469,1539,1565]" collectionCode="NHMUK" pageId="91" pageNumber="92" specimenCode="NHMUK R7994">NHMUK R7994</materialsCitation>
|
||
,
|
||
<materialsCitation collectionCode="CM" pageId="91" pageNumber="92" specimenCode="CM 9380">CM 9380</materialsCitation>
|
||
,
|
||
<materialsCitation box="[598,759,1579,1605]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 23844">LACM 23844</materialsCitation>
|
||
,
|
||
<materialsCitation box="[772,890,1579,1605]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 008">MOR 008</materialsCitation>
|
||
,
|
||
<materialsCitation box="[903,1022,1579,1605]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1035,1167,1579,1605]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 1131">MOR 1131</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1180,1312,1579,1605]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 2822">MOR 2822</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1325,1536,1578,1605]" collectionCode="NMMNH" pageId="91" pageNumber="92" specimenCode="NMMNH P-3698">NMMNH P-3698</materialsCitation>
|
||
,
|
||
<materialsCitation box="[527,702,1619,1645]" collectionCode="UCMP" pageId="91" pageNumber="92" specimenCode="UCMP 118742">UCMP 118742</materialsCitation>
|
||
,
|
||
<materialsCitation box="[714,884,1619,1645]" collectionCode="UMNH" pageId="91" pageNumber="92" specimenCode="UMNH 11000">UMNH 11000</materialsCitation>
|
||
, and
|
||
<materialsCitation box="[946,1093,1619,1645]" collectionCode="UWGM" pageId="91" pageNumber="92" specimenCode="UWGM 181">UWGM 181</materialsCitation>
|
||
, should appropriate bones be available for analysis. However, an EFS is absent from
|
||
<materialsCitation box="[1064,1180,1658,1685]" collectionCode="MOR" pageId="91" pageNumber="92" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
(
|
||
<bibRefCitation author="Horner JR & Padian K." box="[1198,1475,1658,1685]" journalOrPublisher="Proceedings of the Royal Society of London. Series B: Biological Sciences" pageId="91" pageNumber="92" pagination="1875 - 1880" part="271" refId="ref54442" refString="Horner JR, Padian K. 2004. Age and growth dynamics of Tyrannosaurus rex. Proceedings of the Royal Society of London. Series B: Biological Sciences 271 (1551): 1875 - 1880 DOI 10.1098 / rspb. 2004.2829." title="Age and growth dynamics of Tyrannosaurus rex" type="journal article" year="2004">
|
||
<emphasis box="[1198,1475,1658,1685]" italics="true" pageId="91" pageNumber="92">Horner & Padian, 2004</emphasis>
|
||
</bibRefCitation>
|
||
), which suggests that the congruence between the presence of an EFS, maturity, and chronological age is not tightly constrained. Specific predictions of LAG-based chronological ages for specimens in this study are given in
|
||
<tableCitation box="[1226,1334,1778,1804]" captionStart="Table 24" captionStartId="91.[525,580,248,269]" captionTargetBox="[516,1499,320,846]" captionText="Table 24 Predicted LAG counts for Tyrannosaurus rex specimens extrapolated from the ontogram. Estimated ages are indicated by a tilde (~), histologically aged specimens are in boldface." httpUri="http://table.plazi.org/id/DF5C84E2FF9FFFF3FDEBFF70327DFE97" pageId="91" pageNumber="92" tableUuid="DF5C84E2FF9FFFF3FDEBFF70327DFE97">Table 24.</tableCitation>
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,1539,1823,1934]" box="[498,1485,1823,1850]" pageId="91" pageNumber="92">
|
||
2. EFS will be absent from
|
||
<materialsCitation box="[817,1052,1823,1850]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=47861" pageId="91" pageNumber="92" specimenCode="AMNH FARB 5117">AMNH FARB 5117</materialsCitation>
|
||
,
|
||
<materialsCitation box="[1065,1241,1823,1850]" collectionCode="LACM" pageId="91" pageNumber="92" specimenCode="LACM 150167">LACM 150167</materialsCitation>
|
||
, and
|
||
<materialsCitation box="[1306,1481,1823,1849]" collectionCode="UWBM" pageId="91" pageNumber="92" specimenCode="UWBM 99000">UWBM 99000</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="91.[498,1539,1823,1934]" pageId="91" pageNumber="92">
|
||
3. If senescence in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[721,789,1869,1894]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="91" pageNumber="92" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[721,789,1869,1894]" italics="true" pageId="91" pageNumber="92">T. rex</emphasis>
|
||
</taxonomicName>
|
||
does not imply reproductive nonviability, then medullary bone will be found in pregnant senescent females.
|
||
</paragraph>
|
||
<paragraph blockId="92.[498,1542,234,710]" pageId="92" pageNumber="93">
|
||
4. If the distinct ridges of the subcutaneous surface of the maxilla result from ossification of the dermis, then histological sections will show an overlay of metaplastic bone apposed to the pericortical surface (sensu
|
||
<bibRefCitation author="Horner JR & Goodwin MB" box="[1023,1322,314,340]" journalOrPublisher="PLOS ONE" pageId="92" pageNumber="93" pagination="e 7626" part="4" refId="ref54525" refString="Horner JR, Goodwin MB. 2009. Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus. PLOS ONE 4 (10): e 7626 DOI 10.1371 / journal. pone. 0007626." title="Extreme cranial ontogeny in the Upper Cretaceous dinosaur Pachycephalosaurus" type="journal article" year="2009">
|
||
<emphasis box="[1023,1322,314,340]" italics="true" pageId="92" pageNumber="93">Horner & Goodwin, 2009</emphasis>
|
||
</bibRefCitation>
|
||
). That is, if flat facial scales precede the onset of armor-like skin and ossification of the dermis, then histological analysis will show a transition from the absence of metaplastic bone to its presence.
|
||
</paragraph>
|
||
<paragraph blockId="92.[498,1542,234,710]" pageId="92" pageNumber="93">5. If secondary metamorphosis is present, the relationship between bite force and independent measures of body size will be nonlinear.</paragraph>
|
||
<paragraph blockId="92.[498,1542,234,710]" pageId="92" pageNumber="93">6. In terms of quantitative functional morphology, complete subadult skulls will be found to be more similar to adults than to juveniles, indicating an abrupt and wholesale morphological transition from low to tall skulls, whereas the postcranium will show a continuum from juvenile to adult.</paragraph>
|
||
<paragraph blockId="92.[498,770,773,808]" box="[498,770,773,808]" pageId="92" pageNumber="93">
|
||
<heading allCaps="true" bold="true" box="[498,770,773,808]" fontSize="14" level="1" pageId="92" pageNumber="93" reason="0">
|
||
<emphasis bold="true" box="[498,770,773,808]" pageId="92" pageNumber="93">CONCLUSIONS</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="92.[512,1542,842,994]" pageId="92" pageNumber="93">
|
||
1. A single growth series for
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[838,904,843,868]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="92" pageNumber="93" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[838,904,843,868]" italics="true" pageId="92" pageNumber="93">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was obtained that can be divided into 21 growth stages;
|
||
<materialsCitation box="[540,725,883,909]" collectionCode="FMNH" pageId="92" pageNumber="93" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
was recovered as the most mature specimen, whereas
|
||
<materialsCitation box="[1370,1512,882,909]" collectionCode="RSM" pageId="92" pageNumber="93" specimenCode="RSM 2523.8">RSM 2523.8</materialsCitation>
|
||
is one of the least mature adults.
|
||
</paragraph>
|
||
<paragraph blockId="92.[512,1542,842,994]" box="[512,1542,967,994]" pageId="92" pageNumber="93">2. Specimens coded with as few as 1.8% of the characters are in the backbone ontogram.</paragraph>
|
||
<paragraph blockId="92.[512,1541,1013,1413]" pageId="92" pageNumber="93">3. Specimen completeness does not influence the number of unambiguously-optimized synontomorphies that support each node.</paragraph>
|
||
<paragraph blockId="92.[512,1541,1013,1413]" pageId="92" pageNumber="93">4. Five growth categories (juvenile, subadult, young adult, adult, senescent adult) were diagnosed based on histology, synontomorphies, and, in part, size and mass; sharp boundaries between categories are seen at the subadult and adult categories.</paragraph>
|
||
<paragraph blockId="92.[512,1541,1013,1413]" pageId="92" pageNumber="93">5. The sample is numerically biased toward specimens in the subadult, young adult, adult, and senescent adult categories; that is, there is a large gap in the growth series at the juvenile to subadult transition. Also, juveniles are underrepresented in the sample.</paragraph>
|
||
<paragraph blockId="92.[512,1541,1013,1413]" pageId="92" pageNumber="93">
|
||
6. The problematic specimen
|
||
<materialsCitation box="[861,1034,1347,1374]" collectionCode="TMM" pageId="92" pageNumber="93" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
|
||
is a subadult, which accounts for its differences from adults.
|
||
</paragraph>
|
||
<paragraph blockId="92.[512,1527,1432,1538]" pageId="92" pageNumber="93">
|
||
7. Phylogenetic and nonphylogenetic changes follow a similar frequency distribution; phylogenetic changes are more frequent early in ontogeny (growth stages 5
|
||
<emphasis box="[1427,1442,1473,1498]" italics="true" pageId="92" pageNumber="93">–</emphasis>
|
||
9) than later in growth.
|
||
</paragraph>
|
||
<paragraph blockId="92.[498,1517,1557,1759]" box="[512,1231,1557,1584]" pageId="92" pageNumber="93">8. Mandibular changes are completed before cranial changes.</paragraph>
|
||
<paragraph blockId="92.[498,1517,1557,1759]" box="[512,1479,1602,1629]" pageId="92" pageNumber="93">9. The dorsotemporal fossa of the frontal was an origin for adductor musculature.</paragraph>
|
||
<paragraph blockId="92.[498,1517,1557,1759]" pageId="92" pageNumber="93">10. The greatest number of growth changes are seen in the large juvenile and subadult growth categories, at growth stages 5 and 6.</paragraph>
|
||
<paragraph blockId="92.[498,1517,1557,1759]" box="[498,1364,1732,1759]" pageId="92" pageNumber="93">11. The number of growth changes generally decreases during adulthood.</paragraph>
|
||
<paragraph blockId="92.[498,1233,1778,1804]" box="[498,1233,1778,1804]" pageId="92" pageNumber="93">12. The skull roof module undergoes the most growth change.</paragraph>
|
||
<paragraph blockId="92.[498,1530,1823,1895]" box="[498,1485,1823,1850]" pageId="92" pageNumber="93">13. Pneumatic changes are dominated by the antorbital (=paranasal) air sac system.</paragraph>
|
||
<paragraph blockId="92.[498,1530,1823,1895]" box="[498,1530,1868,1895]" pageId="92" pageNumber="93">14. The skull frame is dominant among apneumatic changes in the skull and mandible.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" pageId="93" pageNumber="94">15. Decreases in maxillary and dentary tooth count are broadly congruent with maturity; tooth count in both bones initially increases before it decreases.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" box="[498,1542,319,346]" pageId="93" pageNumber="94">16. Most braincase sutures close during the young adult category, whereas two stay open.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" pageId="93" pageNumber="94">17. Changes to the pectoral girdle and pes are dominant early in ontogeny, whereas axial and pelvic girdle changes occur late; the fibula changes throughout. The most postcranial changes happen to the fibula.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" box="[498,1383,489,516]" pageId="93" pageNumber="94">18. Body size and mass are not congruent with maturity during adulthood.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" pageId="93" pageNumber="94">19. Maturity is congruent with chronological age, bite force, REQ, the presence and depth of the dentary groove, and decrease in agility.</paragraph>
|
||
<paragraph blockId="93.[498,1542,234,685]" pageId="93" pageNumber="94">20. Maturity is incongruent with geographic location, stratigraphic position, phylogeny, and ischial divergence.</paragraph>
|
||
<paragraph blockId="93.[498,1542,704,1100]" pageId="93" pageNumber="94">21. The transition from a long and low skull to a stout and deep skull (i.e., the juvenile-subadult transition) occurred rapidly within a 2-year time span. Ergo, the osteological correlates of skull strength occurred before adult size (i.e., somatic maturity) was reached.</paragraph>
|
||
<paragraph blockId="93.[498,1542,704,1100]" pageId="93" pageNumber="94">
|
||
22.
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[541,608,870,895]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[541,608,870,895]" italics="true" pageId="93" pageNumber="94">T. rex</emphasis>
|
||
</taxonomicName>
|
||
exceeded the plesiomorphic size and mass of
|
||
<taxonomicName box="[1146,1344,869,895]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
between its 15th and 18th years.
|
||
</paragraph>
|
||
<paragraph blockId="93.[498,1542,704,1100]" pageId="93" pageNumber="94">
|
||
23. Sexual maturity in
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[766,835,955,980]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[766,835,955,980]" italics="true" pageId="93" pageNumber="94">T. rex</emphasis>
|
||
</taxonomicName>
|
||
occurred before its 15th year and as early as its 13th year. Ergo, as in crocodylians and other reptiles, including non-avian dinosaurs (
|
||
<bibRefCitation author="Erickson GM & Rogers KC & Varricchio DJ & Norell MA & Xu X." journalOrPublisher="Biology Letters" pageId="93" pageNumber="94" pagination="558 - 561" part="3" refId="ref53283" refString="Erickson GM, Rogers KC, Varricchio DJ, Norell MA, Xu X. 2007. Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition. Biology Letters 3 (5): 558 - 561 DOI 10.1098 / rsbl. 2007.0254." title="Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition" type="journal article" year="2007">
|
||
<emphasis italics="true" pageId="93" pageNumber="94">Erickson et al., 2007</emphasis>
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Lee AH & Werning S." box="[683,933,1034,1060]" journalOrPublisher="Proceedings of the National Association of Science" pageId="93" pageNumber="94" pagination="582 - 587" part="105" refId="ref54829" refString="Lee AH, Werning S. 2008. Sexual maturity in growing dinosaurs does not fit reptilian growth models. Proceedings of the National Association of Science 105 (2): 582 - 587 DOI 10.1073 / pnas. 0708903105." title="Sexual maturity in growing dinosaurs does not fit reptilian growth models" type="journal article" year="2008">
|
||
<emphasis box="[683,933,1034,1060]" italics="true" pageId="93" pageNumber="94">Lee & Werning, 2008</emphasis>
|
||
</bibRefCitation>
|
||
),
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[958,1027,1035,1060]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[958,1027,1035,1060]" italics="true" pageId="93" pageNumber="94">T. rex</emphasis>
|
||
</taxonomicName>
|
||
reached sexual maturity before reaching asymptotic adult size.
|
||
</paragraph>
|
||
<paragraph blockId="93.[498,1190,1119,1145]" box="[498,1190,1119,1145]" pageId="93" pageNumber="94">
|
||
24. Skeletodental sexual dimorphism is absent from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1115,1184,1120,1145]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1115,1184,1120,1145]" italics="true" pageId="93" pageNumber="94">T. rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="93.[498,1542,1164,1360]" box="[498,1490,1164,1191]" pageId="93" pageNumber="94">
|
||
25. There is no evidence for the taxon that is informally dubbed
|
||
<emphasis box="[1263,1485,1164,1190]" italics="true" pageId="93" pageNumber="94">
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1263,1438,1165,1190]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
“x”
|
||
</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="93.[498,1542,1164,1360]" pageId="93" pageNumber="94">26. The entire skeleton is affected by individual variation; reversals of individual variation did not occur until young adulthood (growth stage 8).</paragraph>
|
||
<paragraph blockId="93.[498,1542,1164,1360]" pageId="93" pageNumber="94">27. Individual variation affects the snout dorsum the most; display features are not disproportionately affected by individual variation.</paragraph>
|
||
<paragraph blockId="93.[498,1508,1379,1536]" pageId="93" pageNumber="94">
|
||
28. The variability in adult size implies that
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1019,1088,1380,1405]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="93" pageNumber="94" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1019,1088,1380,1405]" italics="true" pageId="93" pageNumber="94">T. rex</emphasis>
|
||
</taxonomicName>
|
||
had determinate growth type II (sensu
|
||
<bibRefCitation author="Sebens KP" box="[624,770,1419,1445]" journalOrPublisher="Annual Review of Ecology and Systematics" pageId="93" pageNumber="94" pagination="371 - 407" part="18" refId="ref56012" refString="Sebens KP. 1987. The ecology of indeterminate growth in animals. Annual Review of Ecology and Systematics 18 (1): 371 - 407 DOI 10.1146 / annurev. es. 18.110187.002103." title="The ecology of indeterminate growth in animals" type="journal article" year="1987">
|
||
<emphasis box="[624,770,1419,1445]" italics="true" pageId="93" pageNumber="94">Sebens, 1987</emphasis>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="93.[498,1508,1379,1536]" box="[498,1508,1464,1491]" pageId="93" pageNumber="94">29. Growth patterns of the skull do not reflect the distribution of stress loads upon it.</paragraph>
|
||
<paragraph blockId="93.[498,1508,1379,1536]" box="[498,1479,1509,1536]" pageId="93" pageNumber="94">30. The adductor chamber rostrocaudally expands at the young adult growth stage.</paragraph>
|
||
<paragraph blockId="93.[498,1532,1555,1660]" pageId="93" pageNumber="94">31. Removal of redundant putative oversplit characters results in loss of topological resolution, which is almost certainly an indication of the patchiness of the character matrix.</paragraph>
|
||
<paragraph blockId="93.[498,1514,1679,1746]" pageId="93" pageNumber="94">
|
||
32. The type specimen is an adult; three of the characters of Osborn
|
||
<emphasis box="[1302,1308,1679,1705]" italics="true" pageId="93" pageNumber="94">’</emphasis>
|
||
s diagnosis of the taxon are defensible.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |